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Published quarterly by The Lepidopterists’ Society 


THE LEPIDOPTERISTS’ SOCIETY 


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JOURNAL OF 


Tue LepiporreristTs’ SOCIETY 


Volume 56 


2002 


Number | 


Journal of the Lepidopterists’ Society 


56(1), 2002, 14 


LIFE HISTORY AND IMMATURE STAGES OF CATOCALA ATOCALA (NOCTUIDAE) 


LAWRENCE F. GALL 
Entomology Division, Peabody Museum of Natural History, Yale University, New Haven, Connecticut 06520, USA 


JOHN W. PEACOCK 
185 Benzler Lust Road, Marion, Ohio 43302, USA 


AND 


JEFFREY R. SLOTTEN 
5421 N.W. 69th Lane, Gainesville, Florida 32653, USA 


ABSTRACT. The immature stages of Catocala atocala are described and illustrated for the first time. Notes on the biology of the adults 


and larval foodplants are provided. 
Additional key words: 


Catocala atocala Brou (1985) was recognized only 
recently as a separate species, having been treated in 
the literature as a variant of C. agrippina Strecker ever 
since Strecker (1874) provided the first illustration. 
Such late recognition of a new Nearctic underwing 
species is unusual given the longstanding interest in 
the genus, and the existence of several dozen 19th and 
early 20th century specimens of both sexes of C. ato- 
cala in institutional collections in North America and 
Europe (Brou 1985:889 mistakenly believed he had 
collected the first known females in the 1970's). Ironi- 
cally, more than 50 years ago the Palearctic lepi- 
dopterist E. A. Dadd understood that C. atocala was a 
separate species, as he prominently labeled a male and 
female in a series of five C. atocala ex Boll collection at 
the Museum fiir Naturkunde (Berlin) as the types of 
“Catabapta torfrida Dadd.” However, to our knowl- 
edge, the name was never published. 

We are not aware of any previously published infor- 
mation on the life history of Catocala atocala other 
than Gall’s (1991) speculation that, based on the exter- 
nal morphology and wing pattern of the adults, the 
foodplant would likely prove to be “pecan hickories” 
(Carya Nutt. Section Apocarya DC.) rather than 
“hickories” sensu strictu (Carya Section Eucarya DC.) 


underwing moths, Louisiana, Mississippi, nutmeg hickory, Carya (Section Apocarya) myristicaeformis. 


(Juglandaceae). Here we report on the successful rear- 
ing of C. atocala from ova deposited by wild-caught fe- 
males, present field observations of adults in Louisiana 
and Mississippi, and suggest a probable natural larval 
foodplant for this species. 


DISTRIBUTIONAL AND BIOLOGICAL NOTES 


In June 1999, LFG and JWP traveled to Louisiana 
and Mississippi to search for C. atocala and elucidate 
its biology. At the time, the only lepidopterist with sig- 
nificant firsthand knowledge of the species was its de- 
scriber, Vernon Brou. His type series of 35 specimens 
was collected between 1975 and 1983. Among those 
specimens, 32 were taken at Edgard, St. John the Bap- 
tist Parish, Louisiana, two were from Weyanoke, West 
Feliciana Parish, Louisiana, and one was from Missis- 
sippi State, Oktibbeha County, Mississippi. Brou (in. 
litt., June 1999) described the type locality as follows: 
“The Edgard . . . location I lived on and collected for 
about 14 years is not easily accessible. I. . . haven't 
been back there for about 15 years, and I’m sure it is 
nearly impossible [now] to get to the pecan tree area. . . 
this area is primarily agriculture (sugar cane), private 
property, or mostly impenetrable swamp requiring a 
pirogue (boat).” The few specimens of C. atocala cap- 


bo 


tured in Tennessee, Arkansas, Illinois, and Oklahoma 
since the 1985 description all came from habitats ap- 
parently comparable to the type locality—low-lying re- 
gions within approximately 30 km of either the Missis- 
sippi River or a major river that drains into it (Brou’s 
Weyanoke locality was nominally in upland woodlands, 
but nevertheless, quite close to the Mississippi River). 

Based on this information, we opted to begin 
searching in the broad vicinity of Weyanoke, Loui- 
siana, making transects using local roads that ran 
largely perpendicular to the Mississippi River, on the 
assumption that the moth’s primary habitat was not 
upland woodlands per se. On the first transect run on 
28 June 1999, JWP found a number of adult C. atocala 
just east of the Mississippi River near Saint Fran- 
cisville, Louisiana. Our further exploration of the Saint 
Francisville area on 29-30 June 1999 revealed that 
adult C. atocala appeared to occur only in shaded 
woodlands in and around the floodplains immediately 
adjacent to the Mississippi River. We successfully ap- 
plied this search protocol on 1—2 July 1999 as we drove 
north along the western border of Mississippi, ulti- 
mately finding adult C. atocala in the following six 
parishes and counties: off State Route 10, 1 km S of 
Saint Francisville, West Feliciana Parish, Louisiana; 
off Carthage Point Road, 4 km W of Carthage, Adams 
County, Mississippi; 4 km NW of Magna Vista, Is- 
saquena County, Mississippi; 5.5 km W of Deeson, Bo- 
livar County, Mississippi; 8 km WSW of Friar’s Point, 
Coahoma County, Mississippi; west of Sam’s Town 
Casino, 12.5 km W of Robinsonville, Tunica County, 
Mississippi. Despite an extended search on 2 July 
1999, we failed to locate adult C. atocala W of Walls in 
DeSoto County, Mississippi (in that area, on the out- 
skirts of Memphis, Tennessee, the appropriate habitat 
appeared significantly more fragmented and less dense 
than in all counties to the south; but note the prior 
capture of C. atocala by M. Furr in Meeman Shelby 
State Forest, Shelby County, Tennessee). On 3-5 July 
1999, JRS visited the Saint Francisville locality and 
made further observations on C. atocala adults. 

The preferred habitat of C. atocala in Louisiana and 
Mississippi appears to be shaded, moist woodlands in 
lowland floodplains adjacent to the Mississippi River 
with concentrations of box elder (Acer negundo L.) 
(Aceraceae), sycamore (Platanus occidentalis L.) (Pla- 
tanaceae), sassafrass (Sassafrass albidum (Nutt.) Nees) 
(Lauraceae), nutmeg hickory (Carya Section Apo- 
carya myristicaeformis (Michx. f. Nutt.) (Juglan- 
daceae), water locust (Gleditsia aquatica Marsh.) 
(Fabaceae), and various oaks (Quercus sp.) (Fa- 
gaceae). All our adult C. atocala were taken by flush- 
ing them from resting sites on tree trunks during day- 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


light hours—none were attracted to our bait traps at 
night (deployed only one night), and we did not em- 
ploy MV or UV light traps. Adult C. atocala were com- 
mon at the West Feliciana Parish, Adams County, and 
Issaquena County sites, with up to 10-20 moths per 
hour observed at each. Adults were seen resting head 
down from 0.5 to 2.0 m above the ground on the 
trunks of various trees, most often box elder and nut- 
meg hickory, and usually adjacent to or under vines ca. 
1-4 cm in diameter. When startled, adult C. atocala 
proved to be among the wariest of any Catocala 
species we have encountered, and displayed a pen- 
chant for alighting repeatedly on the often inaccessible 
vines; pursuits of 10-20 minutes per moth were not 
uncommon. Adult C. atocala were flushed from trees 
as early as 0715 h at the Saint Francisville locality. 
Daytime high temperatures throughout the period 28 
June to 2 July 1999 ranged from 31—35°C. 

The only other Catocala species occurring in signif- 
icant numbers with C. atocala in these wooded flood- 
plains was C. agrippina. In the Adams County and (es- 
pecially) Issaquena County localities, adult C. atocala 
outnumbered adult C. agrippina. The Catocala 
species encountered at each locality were as follows: 
Louisiana, West Feliciana Parish: C. amatrix Hiibner, 
C. agrippina, C. amica Hiibner, C. atocala, C. caris- 
sima Hulst, C. ilia Cramer, C. insolabilis Guenée, C. 
maestosa Hulst, C. piatrix Grote, C. vidua J. E. Smith; 
Mississippi, Adams County: C. agrippina, C. atocala, 
C. amatrix, C. carissima, C. innubens Guenée; Missis- 
sippi, Bolivar County: C. atocala, C. agrippina, C. in- 
nubens; Mississippi, Coahoma County: C. agrippina, 
C. atocala, C. carissima, C. innubens: Mississippi, Tu- 
nica County: C. agrippina, C. atocala, C. innubens; 
Mississippi, DeSoto County: C. maestosa. 


REARING OBSERVATIONS 


Ova were secured from two of six moderately worn 
female C. atocala collected during the day from Saint 
Francisville, Louisiana, between 30 June and 4 July 
1999. The females were confined in paper grocery 
bags (8 x 13.5 x 27 cm) at near ambient temperature 
(15-30°C), and were offered a weak sucrose solution 
daily on small pieces of sponge; a new sponge piece 
with solution was provided approximately every other 
day. Although females lived from three to eight days, 
only two oviposited, and each of these deposited only 
three ova. One set of ova was overwintered at ambient 
outside temperatures in Gainesville, Alachua County, 
Florida by JRS (coldest periods of circa 5 to 10°C), and 
another in a protected garage in Westport, Fairfield 
County, Connecticut by LFG (coldest periods of circa 
—2 to 5°C). Overwintering ova were kept in empty 


VOLUME 56, NUMBER 1 3 


Fic. 1. Immature stages and distribution of Catocala atocala. a, lateral view of 6th (last) instar larva. b, closeup of head capsule of 6th in- 
star larva. ec, dorsal view of egg as revealed by scanning electron microscopy (60x). d, distribution of Catocala atocala in North America; open 
circles, pre-1950 records; filled circles, post-1950 records; each circle represents a county in which the species has been recorded; shading rep- 
resents distribution of Carya (Section Apocarya) myristicaeformis (after Little 1977, 1980). 


plastic film containers and misted lightly once every one emerged on 12 May 2000 from the three ova over- 
few weeks. wintered in Connecticut. The Florida larva was of- 
Only one first instar larva emerged on 14 April 2000 fered only water hickory (Carya Section Apocarya 


from the three ova overwintered in Florida, and only aquatica (Michx. f.) Nutt.), which it accepied. The 


Connecticut larva was offered an array of potential 
jungladaceous foodplants in an arena food choice test 
(see Gall 1991); it accepted black walnut (Juglans ni- 
gra L.) and butternut (J. cinerea L.), but refused to 
feed on shagbark hickory (C. Section Eucarya ovata 
(Mill.) K. Koch), pignut hickory (C. Section Eucarya 
glabra (Mill.) Sweet), and bitternut hickory (C. Sec- 
tion Apocarya cordiformis (Wang.) K. Koch). This 
larva subsequently was reared on J. nigra. Both larvae 
were reared indoors at 20-24°C, and each appeared to 
eat their respective foods equally well, as development 
times per larval instar differed by one day or less. Both 
larvae were preserved in alcohol as fully grown last in- 
stars. The average duration for each stage was as fol- 
lows (based on n = 2 larvae): Ist instar, 4 days; 2nd in- 
star, 4 days; 3rd instar, 3.5 days; 4th instar, 4.5 days; Sth 
instar, 5.5 days; 6th instar >5 days (larvae were pre- 
served before entering pre-pupal stage). A description 


of the mature larva is as follows: 

6th (final) instar larva (Figs. la, b). Length 6 cm; head capsule 
width 3-4 mm; body color light mousy grey tinged with pink; dorsal 
tubercles reddish orange; spiracles black; finely dispersed black to 
brown specks forming interrupted but nearly complete dorsal and 
lateral lines along length of body, most apparent in the vicinty of the 
dorsal tubercles; 5th abdominal segment with slightly elevated, 2 
mm wide transverse protuberance, slightly lighter than body color, 
with a large dark brown lateral “saddle patch” contrasting strongly 
with the body color, all lines formed by specks being darker and 
more prominent on this segment; 8th abdominal segment with a 
pair of ventrally projecting, 1 mm long tubercles; lateral filaments 
present along entire length of body, whitish pink, dense, 1-2 mm in 
length, simple (not bifurcate or multifurcate); ventral surface of 
body segments dirty whitish pink, with black spots on each abdomi- 
nal segment, edged and overlayed with orange on 4th through 7th 
abdominal segments; capitad surface of head capsule flattened but 
not strongly produced addorsally, lacking lateral stripes from anten- 
nae to dorsal margins; antennae and true legs pinkish; setae on head 
capsule and body pinkish, sparse; head capsule color same as body 
color, but with slightly darker striations; based on limited sample 
sizes, the larva of C. atocala appears to have a more prominent “sad- 
dle patch” than the larva of C. agrippina, and the first few dorsal tu- 
bercles behind the head capsule in C. agrippina are usually yellow- 
ish, as opposed to reddish orange in C. atocala. 


It is unlikely that Carya aquatica or Juglans nigra 
are the wild larval foodplants of C. atocala at the 
Louisiana or Mississippi localities visited during 1999. 
No Carya aquatica were seen at these localities, and J. 
nigra, Carya Section Eucarya ovata, and Carya Section 
Apocarya illinoiensis (Wang.) K. Koch (pecan) occurred 
only sporadically, and then not in close proximity to the 
adult moths. However, Carya myristicaeformis, the only 
other juglandaceous tree species at any of the C. atocala 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


collecting sites, was present and often common where 
the moths were found, and was one of the tree species 
on which the adults frequently rested during the day- 
time. In addition, the overall geographic range of 
Carya myristicaeformis approximates the known 
range of C. atocala (Fig. 1d). We thus believe Carya 
myristicaeformis is the wild foodplant for C. atocala 
along the Mississippi River in Louisiana and Missis- 
sippi. Because another global stronghold for Carya 
myristicaeformis appears to be along the Red River 
and its tributaries on the border of Oklahoma and 
Texas, we predict that C. atocala occurs there in 
greater numbers than the few recent Oklahoma cap- 
tures (Cleveland, Murray and Tulsa Counties; G. Gier 
& C. Harp, pers. com.) suggest. In addition, disjunct 
populations of Carya myristicaeformis that could har- 
bor C. atocala exist in coastal South Carolina. We en- 
courage lepidopterists to look for C. atocala and Carya 
myristicaeformis in these other drainage systems, as 
well as at greater distances from rivers per se, as our 
sampling efforts during 1999 were at best inconsistent 
the farther we were from the Mississippi River. 


ACKNOWLEDGMENTS 


We thank Vernon Brou for his hospitality in Louisiana during 
1999, and for sharing his catocalating experiences in that state. Wol- 
fram Mey and Wolfgang Speidel provided access to the collections 
at the Museum fiir Naturkunde in Berlin to LFG. Wayne Miller and 
Jim Tuttle helped JRS in the field at the Saint Francisville locality. 
Jeff Cott took the larval photographs, and John Brown provided 
helpful comments on the manuscript. 


LITERATURE CITED 


Brou, V. A. 1985. A new species of Catocala (Lepidoptera: Noctu- 
idae) from the Gulf South, U.S.A. Proc. Entomol. Soc. Wash. 
87:889-892. 

GatL, L. F. 1991. Evolutionary ecology of sympatric Catocala 
moths (Lepidoptera:Noctuidae). I. Experiments on larval food- 
plant specificity. J. Res. Lepid. 29:173-194. 

LITTLE, E. L. 1977. Atlas of United States trees. Vol. 4. Minor east- 
em hardwoods. U.S. Dept. Agric. Misc. Publ. No. 1342. Gov- 
ernment Printing Office, Washington, D.C. 

. 1980. The Audubon Society field guide to North American 
trees. Chanticleer Press, New York, New York. 714 pp. 

STRECKER, F. H. H. 1874. Lepidoptera, Rhophaloceres and Het- 
eroceres indigenous and exotic; with descriptions and colored 
illustrations. No. 11. Owen’s Steam Book and Job Printing Of- 
fice, Reading, Pennsylvania. Pp. 95-100. 


Received for publication 6 January 2001; revised and accepted 22 
November 2001. 


Journdl of the Lepidopterists’ Society 
56(1), 2002, 5-8 


DISTRIBUTION AND BIOLOGY OF ANISOTA MANITOBENSIS (SATURNIIDAE) IN 
SOUTHERN MANITOBA 


DONALD C. HENNE 
Department of Entomology, 402 Life Sciences Building, Louisiana State University, Baton Rouge, Louisiana 70803, USA 


ABSTRACT. The known distribution of the saturniid moth, Anisota manitobensis is limited to southern Manitoba and northern Minnesota, 
To date, nothing has been reported in the literature about adult emergence times and adult mating activity. In this paper, the known distribu- 
tion of A. manitobensis in southern Manitoba is summarized, information on larval collection techniques is provided, and new information on 
adult emergence times and adult mating activities are given. Adults begin emerging at approximately 0600 h (CDT), with a few delaying emer- 
gence until between 2100 and 2200 h (CDT). Adult females of A. manitobensis call from 0630 to 0900 h (CDT) and, if not mated, again from 


0100 to 0300 h (CDT). 


Additional key words: Anisota stigma, bur oak, calling females, larvae, oakworms. 


The saturniid moth Anisota manitobensis McDun- 
nough is known from southern Manitoba, northern 
Minnesota, and may possibly occur in northeastern 
North Dakota (McGugan 1958, Tuskes et al. 1996). 
Anisota manitobensis was reported from Wisconsin by 
Riotte and Peigler (1981). However, these records 
have been disputed by Tuskes et al. (1996). The known 
distribution of A. manitobensis is based largely on very 
old collection records. There are relatively few collec- 
tion records for A. manitobensis, and almost none 
from the last 25 years. Nothing was known about adult 
emergence times, adult female calling times, the 
length of the mating period, or oviposition behavior of 
this species. 

This paper summarizes most of the available collec- 
tion records for this moth in southern Manitoba. 
These locality records were utilized in an attempt to 
locate this species in the field. In 1996, and again in 
1997, A. manitobensis was collected at Fullers, Mani- 
toba. Larvae were located on very small bur oak (Quer- 
cus macrocarpa Michaux) trees, in semi-open areas. 
New information about adult emergence times and fe- 
male calling times indicate that adults begin emerging 
at approximately 0600 h (CDT). A few individuals de- 
layed emergence until between 2100 and 2200 h 
(CDT). Adult females of A. manitobensis call from 
0630 to 0900 h (CDT) and again from 0100 to 0300 h 
(CDT), if not mated during the morning calling period. 


MATERIALS AND METHODS 


A search for colonies of Anisota spp. in southern 
Manitoba was conducted annually from 1989 to 1996. 
Using the available collection records, many of the lo- 
calities in Manitoba where species of this genus have 
been collected in the past were extensively searched, 
often on more than one occasion. With the exception 
of a single locality record for A. virginiensis (Drury) at 
Belair, Manitoba, efforts at locating Anisota spp. (e.g., 
black lighting, searching for larvae) yielded no addi- 
tional colonies. Virgin females of the closely related A. 
stigma Fabricius (courtesy of Mr. J. P. Tuttle) were also 


employed in an effort to locate colonies of A. manito- 
bensis in and around Winnipeg. 

Anisota manitobensis larvae were maintained on fo- 
liage of Q. macrocarpa. Pupae were placed in moist 
peat and overwintered at 5°C. Pupae were removed 
from cold storage in early May and placed in shaded 
outdoor cages to ensure exposure to ambient temper- 
atures and photoperiod. Voucher specimens have been 
retained in the personal collection of the author. 


RESULTS 


Historical distribution (Fig. 1). According to the 
relevant literature (McDunnough 1921, Brodie 1929, 
McGugan 1958, Ferguson 1972, Riotte & Peigler 
1981, Tuskes et al. 1996), and museum records (J. B. 
Wallis Museum of Entomology [JBWM], Manitoba 
Museum of Man and Nature [MMMN], and the 
Transcona Historical Museum [THM]), A. manitoben- 
sis has been collected at the following southern Mani- 
toba localities: 

Anola: 26 June 1976 (Tuskes et al. 1996), Aweme 
(8 km north of Treesbank): 23 June 1904 (McDun- 
nough 1921), 29 June 1907 and 4 July 1907 (McDun- 
nough 1921), ([larva] August 1912 [JBWM}]), Birds 
Hill: 20 July 1963 (THM), Brandon: 15 July 1950 
(JBWM), Darlingford: (Riotte & Peigler 1981), Kel- 
wood: (Riotte & Peigler 1981), Killarney: 10 July 
1947 (J]BWM), McCreary: (Riotte & Peigler 1981), 
Middlechurch: July 1954 (J[BWM), Pine Ridge: 
(Riotte & Peigler 1981), Riding Mountain National 
Park: 19 June 1936 (Ferguson 1971), Sandilands 
Provincial Forest: 25 June 1971 (C. S. Quelch field 
notes—THM), Souris: June 1954 (JBWM), 1 mi. W. 
Vivian: 30 June 1967 (C. S. Quelch field notes— 
THM), Winnipeg: July 1920 (JBWM), 10 June 1921 
and 16 June 1921 (JBWM), 8 June 1928 (]BWM), 29 
June 1928 (Brodie 1929), 3 July 1930 (JBWM), 22 
June 1948 (JBWM), 5 July 1949 (BWM), 10 July 1950 
(THM), 26 June 1954 and 2 July 1954 (THM), 14 July 
1955 (THM). Criddle (1932) reported A. manitobensis 
from the area south of Carman, where three acres of bur 


6 


Fic. 1. Distribution of Anisota manitobensis in southern Mani- 
toba. Locality of most recent collection at Fullers indicated by arrow. 


oak were severely defoliated in 1931. Criddle also men- 
tioned Onah and Treesbank as other localities where 
A. manitobensis larvae were collected that same year. 
New collection locales. On 20 August 1996, two 
late fifth-instar A. manitobensis larvae of opposite sex 
were collected east of Fullers, Manitoba. This locality 
is approximately 5 km north of East Selkirk, along 
highway #508. As with other saturniid larvae, larvae of 
Anisota can be sexed according to the presence or ab- 
sence of a small dark spot on the venter of the ninth 
abdominal segment. This spot is associated with the 
male genital histoblast (Miller 1977). Larvae having this 
spot are males and those lacking this spot are females. 
These larvae were found feeding on the foliage of a 
small bur oak that was approximately 150-180 cm in 
height. An egg cluster containing 16 eggs was located 
on the tip of an oak leaf at the end of a branch, ap- 
proximately 45 cm above ground level and facing 
south. Of these 16 eggs, 12 had eclosed and the other 
four contained dead embryos. On 22 August 1996 a 
second collection consisting of a single late fifth-instar 
female larva was also made, approximately 1 km west 
of the first collection. This larva was also collected 
from a small bur oak tree that was only 75 cm in height 
and >90% defoliated; no egg cluster was located. It 
was evident that several other larvae were also present 
on this tree but had probably wandered away to pu- 
pate. At both collection points, the oak trees on which 
these larvae were collected were isolated (i.e., > 150 
cm) from other trees, had low surrounding vegetation, 
and were receiving full sunlight. In the laboratory, all 
three larvae wandered for approximately two days af- 
ter feeding was completed, and pupated approxi- 
mately three days after wandering ceased. All larvae 
turned a mottled green color several days prior to pu- 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


pation. The same locality was revisited in June 1997. A 
cluster of 75 second-instar larvae was found on a ter- 
minal oak leaf, only 30 cm above the ground and fac- 
ing west, in full sunlight, and along the forest-field in- 
terface. Sixty-five pupae were obtained from this 
collection (32 males, 33 females). 

Morphological comparisons (Fig. 2). According 
to Brodie (1929) mature A. manitobensis larvae pos- 
sess pale tan head capsules. Examination of the larvae 
collected near Fullers revealed that the head capsule is 
clearly orange, as in mature larvae of the closely re- 
lated A. stigma (Riotte & Peigler 1981). In addition, 
the single larva collected on 22 August 1996 differed 
from the other two in that the dorsal stripe and shad- 
ing described by Brodie were very faint and barely 
visible. Color photos of these larvae can be viewed at 
http:/Avww.lsu.edu/faculty/dhennel/. Brodie (1929) 
also described the pupa of A. manitobensis as reddish- 
brown in color. Although this color was observed for 
the first few days following pupation, the pupae even- 
tually darkened to a brownish-black color. Confirma- 
tion of the larval collections as being A. manitobensis 
was obtained the following June when adults emerged 
from over wintered pupae. Adults closely resembled 
museum specimens of A. manitobensis. The speci- 
mens obtained lacked any heavy spotting of the wings, 
a trait characteristic of A. stigma. Examination of male 
genitalia of specimens of A. stigma from Anne Arundle 
Co., Maryland, compared with those of the specimens 
collected at Fullers also confirmed the identity of A. 
manitobensis. 

Mating activity. Adult emergence times, female 
calling times, length of pairing, and oviposition habits 
for A. manitobensis were unknown until this time. 
Adults obtained from these collections emerged in the 
morning, around 0600 h (CDT). Females began call- 
ing as soon as their wings were fully expanded, often 
within 30 minutes of eclosion. Most females had their 
ovipositors everted from approximately 0630 to 0900 h 
(CDT). If unmated during this time these females 
would exhibit a second calling time, from approxi- 
mately 0100 to 0300 h (CDT). These calling times are 
similar to those reported for A. stigma (Tuskes et al. 
1996). Some adults delayed emergence until between 
2100 and 2200 h (CDT), which is shortly after sunset 
in southern Manitoba during early June. Length of 
pairing at night was variable, ranging from one to sev- 
eral hours. Adults mated during the morning hours re- 
mained in copula until dusk, at which time females 
would begin ovipositing. This behavior is very similar 
to that reported for A. stigma in Tuskes et al. (1996). 
Oviposition behavior after separation of adults was dif- 
ficult to observe. Most females placed in paper sacks 


VOLUME 56, NUMBER I 


Fic. 2. Fifth-instar larvae of Anisota manitobensis. (A) lateral view, (B) dorsal view 


after pairing ceased refused to lay eggs. A single female, 
however, deposited 185 eggs inside a paper sack, over a 
three-day period. Egg laying activity was not observed. 


DISCUSSION 


Anisota manitobensis was a very difficult species to 
locate in the field. McGugan (1958) reported A. mani- 
tobensis larvae as locally abundant in some areas of the 
Red River Valley of southern Manitoba but also stated 
that it is generally found in small numbers. This would 
appear to be the case with the population at Fullers, 
Manitoba. Evidence of bur oak defoliation typical of 
feeding by larval Anisota was also noted near East 
Selkirk, and in an area east of Winnipeg, just south of 
Pine Ridge. No larvae or egg clusters were found at 
these localities. Gregarious larvae of Datana ministra 
(Drury) were common in these areas and were noted 
for the similar method by which they defoliated bur 
oak. Larvae of Anisota and Datana consume all of the 
oak leaf, with the exception of the mid vein. Riotte and 
Peigler (1981:113) also found Datana to be common 
and similarly interfered/confused their searching for 
Anisota. It was also noted that agricultural areas now 
isolate many of the localities where A. manitobensis 
was collected in the past. Local extirpations of this in- 
sect have likely occurred on a large scale, but it is be- 
yond the scope of this paper to speculate on the popu- 
lation status of this insect. If it is indeed rare, then it 
needs to be protected. We still know very little, how- 
ever, about population dynamics of most Anisota spp. 
The low populations may only be a cyclic phenome- 
non. Alternatively, it may be possible that this species 
prefers oak stands of a limited age, i.e., older stands of 
oak may not be suitable habitat for A. manitobensis. 

The similarity of calling times of A. manitobensis 
and A. stigma females may support the notion of 
Tuskes et al. (1996) that A. manitobensis is only a clinal 


variant of A. stigma. Unfortunately, there is a huge gap 
in collection records between A. manitobensis in Man- 
itoba and the nearest records for A. stigma in Min- 
nesota and Wisconsin. Material from this region could 
be useful in solving the question as to whether A. man- 
itobensis should be recognized as a full species or not. 
The failure of A. stigma females to attract A. manito- 
bensis males does not necessarily indicate that some 
form of prezygotic isolating mechanism isolates the 
two species. Instead, it may mean that A. manitobensis 
was simply not present in the localities where A. 
stigma females were deployed. Unfortunately, live- 
stock of A. stigma was not available to be tested on a 
known population of A. manitobensis. 


ACKNOWLEDGMENTS 


I would like to thank the following individuals for their helpful 
comments, insights, and suggestions in reviewing this manuscript: 
R. S. Peigler (San Antonio, Texas), J. K. Diehl and R. E. Roughley 
(University of Manitoba), and to the following institutions for allow- 
ing me to record specimen label data: J. B. Wallis Museum of Ento- 
mology (University of Manitoba), Manitoba Museum of Man and 
Nature, and the Transcona Historical Museum. I would also like to 
thank Mr. Derek Bridgehouse for providing specimens of Anisota 
stigma from Maryland and Mr. Helios Hernandez (Manitoba De- 
partment of Natural Resources) for assistance in obtaining necessary 
scientific research and collecting permits to study Anisota in Mani- 
toba Provincial Parks. I am also grateful to Ms. Stacy Clayton (LSU 
Department of Entomology) for assistance in posting the web site. 


LITERATURE CITED 


Bropig, H. J. 1929. Notes on the early stages of Anisota manito- 
bensis (Lepid.). Can. Ent. 61:98-100. 

CRIDDLE, N. 1932. The Canadian Insect Pest Review—Forest and 
Shade Tree Insects 10(4):79. 

FERGUSON, D. C. 1972. Bombycoidea (in part), fasc. 20.2B. In R. B. 
Dominick et al. (eds.), The moths of America north of Mexico. 
Curwen Press, London. 

McDunnoucu, J. H. 1921. A new Anisota species from Manitoba 
(Lepid.). Can. Ent. 53:75. 

McGuean, R. M. 1958. Forest Lepidoptera of Canada recorded 
by the Forest Insect Survey. I. Papilionidae to Arctiidae. Forest 
Biology Div., Canada Dept. of Agric., Publ. 1034. Ottawa. 


MILLER, T. A., W. J. COOPER & J. W. HIGHFILL. 1977. Determina- 
tion of sex in four species of giant silkworm moth larvae (Sat- 
urniidae). J. Lepid. Soc, 31:144-146. 

Riorre, J. C. E. & R. S. PEIGLER. 1980 [1981]. A revision of the 
American genus Anisota (Satumiidae). J. Res. Lepid. 19:101-180. 

TuskEs, P. M., J. P. Turrte & M. M. Couns. 1996. The wild silk 
moths of North America: a natural history of the Saturniidae of 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


the United States and Canada. The Cornell Series in Arthropod 
Biology. Comell University Press, Ithaca. 264 pp. 


Received for publication 11 April 2000; revised and accepted 4 
December 2001. 


Journal of the Lepidopterists’ Society 
56(1), 2002, 9-44 


THE LARGE MOTHS OF GUANA ISLAND, BRITISH VIRGIN ISLANDS: A SURVEY OF EFFICIENT 
COLONIZERS (SPHINGIDAE, NOTODONTIDAE, NOCTUIDAE, ARCTIIDAE, GEOMETRIDAE, 
HYBLAEIDAE, COSSIDAE) 


VITOR O. BECKER 


Research Associate, Departamento de Zoologia, Universidade de Brasilia, P.O. Box 04525, 70919-970 Brasilia, DF, Brazil, and Department of 
Systematic Biology, Smithsonian Institution, Washington, DC 20560-0105, USA 


AND 


ScoTT EF. MILLER 
Department of Systematic Biology, Smithsonian Institution, Washington, DC 20560-0105, USA 


“Good boys go to heaven, but the bad boys go everywhere”* 
Meatloaf 1993 


ABSTRACT. An illustrated and annotated list of large moths of Guana, a 297 ha island located on the north side of the Caribbean island 
of Tortola, British Virgin Islands, is presented. Of the 148 species listed, 98 are distributed throughout the neotropics, 41 throughout the An- 
tilles, with some ranging into Florida, and 9 endemic to the Puerto Rican Bank, two of them described here: Catabenoides lazelli, new 
species, and Perigea gloria, new species. The following new synonyms and new combinations are recognized: Leucania solita Walker, new 
synonym [=L. humidicola Guenée], Kakopoda cincta Smith, new synonym [=K. progenies (Guenée)], Drepanopalpia polycyma Hampson, 
new synonym [=D. lunifera (Butler), new combination], Sphacelodes fusilineatus Walker, revised status, Idaea fernaria (Schaus), new com- 
bination, Ptychopoda curtaria Warren, new synonym [=Idaea minuta (Schaus)]. Pterocypha defensata Walker, revised status, is recognized 
as the senior synonym of P. floridata (Walker), new synonym, reversing a recently published synonymy. A new genus, Catabenoides Poole, new 
genus, type-species: Laphygma vitrina Walker, is described in an appendix, including C. divisa (Herrich-Schiffer), new combination, C. se- 
orsa (Todd), new combination, and C. terens (Walker), new combination, all by Robert W. Poole. The palatability to birds of two species, 
Diphthera festiva and Calidota strigosa, was observed and the species were shown to be distasteful. 


Additional key words: Caribbean, West Indies, biogeography, taxonomy, palatability. 


Guana is a small island on the north side of Tortola Schaus 1940, Wolcott 1951), but most have not been 
in the British Virgin Islands (18°28’N, 64°35’W) (Fig. recorded from the British Virgin Islands due to lack of 
1). While it is small, only 297 ha, and the maximum el- previous sampling. The faunal similarity to Puerto 
evation is 266 m, it supports a relatively rich vegetation Rico is expected, given that the principal islands of the 
and has sustained less damage by feral animals and Virgin Islands (except Saint Croix) lost their connec- 
humans than have many adjacent islands (Lazell tion with each other and with Puerto Rico only about 
1996). It has most of the floristic associations of the 8000 to 10,000 years ago, due to eustatic rise in sea 
larger Virgin Islands, with the notable exception of the level (Heatwole et al. 1981). Only scattered records ex- 
“aridulate rain forest” of Tortola (D’Arcy 1967). De- ist in the literature for moths of the Virgin Islands, 


spite its small size, Guana has a diverse insect fauna with two of the longest lists being Beatty (1948) for St. 
(Davies & Smith 1997). For example, Guana has 31 Croix and Greenwood and Greenwood (1971) for Pe- 
species of butterflies (Becker & Miller 1992), com- ter Island. 

pared to the larger islands of Anegada (3872 ha) We are treating the Lepidoptera of Guana Island in 
(Smith et al. 1991) with 24 species and Tortola (5444 parts. Becker and Miller (1992) reported 31 species of 
ha) with 31 species, and St. Thomas (7660 ha) with 32 butterflies. The present paper reports 148 species of 


species (Miller 1994). Alminas et al. (1994) review the large moths (Macrolepidoptera, including the unre- 
geographical setting of the Virgin Islands. lated Cossidae and Hyblaeidae for convenience), rep- 

This is the first survey of the moth fauna of this is- resented by 1390 specimens. The manuscript was pre- 
land. John F. G. Clarke was on Guana briefly in 1956 pared using the classification of Noctuidae by Poole 
and 1958 (see Schmitt 1959), but was unable to collect (1989), before extensive recent changes in higher clas- 
at lights there. Most of the species reported herein are sification of Lepidoptera (Kristensen 1998, Holloway 
known from Puerto Rico (e.g., Forbes 1930, 1931, et al. 2001). Future papers will treat Pyraloidea and 

Microlepidoptera. 

*We often pejoratively regard widespread species as “weed The moth fauna of Guana is composed primarily of 

species” or “pests”. Parodying the citation above: “Bad” species go species with wide distributions in the New World trop- 


everywhere, “good” species [may] go to Heaven [extinct]. But is it : , ; . 3 P 
fair to consider them bad simply because they are able to get every- ics (Table 1). Of the 148 Speers listed, 50 are endemic 


where? Perhaps “efficient colonizers” is a better term? to the Caribbean Islands, many of them reaching the 


10 


MUSKMELON $§ 
BAY 


Quail-Dove Ghut 
WHITE BAY 


Harris Ghut 
=, 


Pinguin Ghut 


MONKEY 


POINT 


NORTH 
BAY 


Palm Ghut 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 1. Map of Guana Island. Contours are 20 m. Bar, bottom right, is 400 m. Inset shows the position of the Virgin Islands in the Antilles. 


Florida Peninsula, and only nine apparently are re- 
stricted to the Puerto Rican Bank. These proportions, 
however, are not the same for the different families. 
The highest degree of endemism occurs in the 
Geometridae and Arctiidae, whereas the lowest occurs 
in the Sphingidae. This is presumably because of the 
powerful flying capacity of sphingids, while geometrids 
and arctiids are clumsy flyers. 

Holloway and Nielsen (1998, following Ferguson et 
al. 1991) presented a chart of 12 moth genera that are 
widely recorded from remote islands worldwide. Of 
these, nine genera (including 20 species) are present 
on Guana (counting Leucania as Mythimna). It is 
likely that the remaining three genera, especially Agro- 
tis, may be found on Guana in the future. Many of the 
species whose ranges include the southern United 


States (especially Sphingidae) also occur as vagrants in 
the Northeast United States and into Canada (e.g., 
Forbes 1954, 1960). Species recorded from the Gala- 
pagos Islands by Hayes (1975) are noted as indication 
of their dispersal ability (note that none of the 
Geometridae recorded from Galapagos are also known 
from Guana). 


MATERIALS AND METHODS 


The material upon which this list is based was col- 
lected in July 1984 and 1985 (by S. E. Miller & P. M. 
Miller), July 1986 (S. E. Miller & M. G. Pogue), July 
1987 (S. E. Miller & V. O. Becker), July 1988 (S. E. 
Miller & C. O’Connell), October 1989 (V. O. Becker), 
and October-November 1990 (S. E. Miller & T. M. 
Kuklenski). Collections from 1984-1986 are deposited 


VOLUME 56, NUMBER | 


TABLE 1. Geographic range of the species of moths collected at 
Guana Island. 


Number of species 


Puerto 
Family Neotropical* Antilles Rican Bank Total 
Sphingidae 20 y) 0 99, 
Notodontidae 1 0 0 1 
Noctuidae 63 16 3 82 
Arctiidae 6 5 4 15 
Geometridae 6 18 2 26 
Hyblaeidae 1 0 0 1 
Cossidae 1 0 0 il 
Total 98 Al 9 148 


* Includes Cosmopolitan and Pantropical species. 


at the National Museum of Natural History (USNM), 
Washington, those of 1987-1990 are split between first 
author (VOB), Bishop Museum, Hawaii (BPBM), and 
USNM. The largest collections were made by both au- 
thors 9-23 July 1987, with over 2200 specimens repre- 
senting about 300 morphospecies, and by the first au- 
thor in October 1989, with over 2000 specimens 
representing over 350 morphospecies. 

The list of species by family collected during the 
1989 trip (Table 2) gives an impression of the overall 
fauna, especially the diversity of Microlepidoptera, still 
under study. Of the 359 species recorded, 243 species 
(two thirds) are Microlepidoptera and pyraloids, and 
163 (nearly one half) are pyraloids and gelechioids. 

The number of specimens listed in this work under 
each species does not reflect relative abundance, as 
our objective was only to list the species occurring on 
the island. Therefore, in the case of species that are 
common in other parts of the neotropics only one or a 
few specimens were collected to voucher the record, 
independent of their abundance. In contrast, in the 
case of endemic or rare species, usually all specimens 
were caught. 

The results obtained by the first author during the 
1989 trip were a surprise, as it was made 20-40 days 
after Hurricane Hugo had devastated the island on 18 
September. The damage was still evident: all buildings 
without roofs and without most of their doors and win- 
dows, and fallen trees and torn branches scattered all 
over the island. According to people who were on the 
island the day Hugo hit, not a single leaf remained on 
the trees. At the time of arrival (9 October) the island 
was completely green again. That same night the col- 
lecting was fantastic, as were most nights during the 
next 20 days. Not only the quantity, but also the qual- 
ity of the material was impressive. It seemed that all 
specimens had emerged that day. Certainly all cater- 
pillars that had reached development had to pupate 


TABLE 2. Moth species collected on Guana Island between 10 
and 20 October 1989. 


Family Number of species 
Sphingidae 10 
Notodontidae ] 
Noctuidae 69 
Arctiidae 12 
Geometridae 24 
Hyblaeidae 1 
Crambidae 51 
Pyralidae 36 
Pterophoridae 4 
Oecophoridae 2 
Blastobasidae 13 
Gelechiidae 45 
Scythrididae 1 
Cosmopterigidae 15 
Psychidae 1 
Tineidae 35 
Gracillariidae 13 
Yponomeutidae 1 
Argyresthiidae 1 
Heliodinidae 2 
Choreutidae 1 
Cossidae 1 
Tortricidae 19 
Opostegidae 1 
Total 359 


immediately after the hurricane, and they were all 
emerging together. Leaf mines were also abundant. As 
these tiny species usually have shorter life cycles, they 
had time to emerge and lay eggs in the 20 days that 
preceded the field work. Torres (1992) documented 
the impact of Hurricane Hugo on Lepidoptera popu- 
lations on Puerto Rico. 

A synoptic collection, containing at least one speci- 
men representing each form, was taken to the Smith- 
sonian Institution (USNM), Cornell University (CU), 
and most importantly, to the Natural History Museum 
(BMNH), London, by the first author. The list below is 
a result of the identifications made by comparing this 
synoptic collection with identified material, especially 
type specimens (including important voucher speci- 
mens and types from Forbes 1930, 1931, Schaus 
1940). The first author has compiled a synonymic list 
of Antillean Lepidoptera (Becker in prep.), which has 
provided further taxonomic background. 

This report is aimed not only at lepidopterists, but 
also at biologists and students interested in the fauna 
of the Virgin Islands. For this reason we give a brief 
synopsis of each species to provide a context and to 
suggest what kind of observations should be under- 
taken. Information on each species is provided under 
the following headings: ID: Diagnostic characters al- 
lowing identification of the species in the context of 


12 


the other species know from Guana Island; DIST: 
General distribution of the species, as represented in 
the literature and VOB and USNM collections; 
GUANA: The Guana Island specimens seen by us in 
preparing the manuscript (see the introduction for the 
dates sampled each year); BIO: Known host records, 
sometimes including notes on immature stages; COM: 
Any other comments. 


SPECIES ACCOUNTS 


SPHINGIDAE 


Eighty five species of sphingids have been recorded 
from the Antilles (Becker in prep.), 22 (25%) of them 
were collected by us in Guana. Color illustrations of 
adults can be found in Hodges (1971) and D’Abrera 
(1986): color illustrations of larvae can be found in 
Moss (1912, 1920). 


Sphinginae 


Agrius cingulatus (Fabricius, 1775) 
(Sweetpotato hornworm, Pink spotted hawk moth) 


ID: Separated from other hawk moths by the pink dots 
on the abdomen. 

DIST: United States south to Argentina, including 
Galapagos. 

GUANA: | specimen, 1989. 

BIO: The variable larvae (green to dark brown) feed 
on various plants belonging to Convolvulaceae, includ- 
ing sweet potato. 


Cocytius antaeus (Drury, 1773) 
(Giant sphinx) 


ID: Distinguished from other sphingids on the island 
by the dark green wings and three pairs of yellow dots 
on the abdomen. 

DIST: Southern United States to Argentina; not in 
Galapagos. 

GUANA: | specimen, 1989. 

BIO: Green larvae feed on various Annona species 
(Annonaceae) such as custard apple. 


Manduca sexta (Linnaeus, 1763) 
(Tobacco hornworm, Carolina sphinx) 


ID: Distinguished from other sphingids on the island 
by the gray wings and six pairs of yellow dots on the 
abdomen. 

DIST: Widespread in the New World, including Gala- 
pagos. 

GUANA: 5 specimens, 1984, 1988, 1989. 

BIO: Pest of cultivated solanaceous plants, such as to- 
bacco, tomato, potato, etc. Mature larvae are green- 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


yellow with seven pairs of white lateral bands, and red 
anal horn. 


Manduca rustica (Fabricius, 1775) 
(Rustic sphinx) 


ID: Distinguished from other sphinx moths on the is- 
land by the dark grayish-brown forewing with trans- 
verse grayish-white waving bands, and three pairs of 
yellow dots on the abdomen. 

DIST: Widespread New World species, present in 
Galapagos. 

GUANA: 2 specimens, 1984, 1989. 

BIO: Larva is greenish-gray, distinguished by its small 
white nodules on the thoracic segments, mainly on 
dorsum. They feed on various species of Bignoniaceae, 
Verbenaceae and Boraginaceae (Hodges 1971). 


Manduca brontes (Drury, 1773) 


ID: Distinguished from other species in the genus oc- 
curring on the island by the absence of yellow dots on 
the abdomen. 

DIST: Antillean species occurring northward to Cen- 
tral Florida. 

GUANA: | specimen, 1989. 

BIO: The larva is similar to that of the tobacco hornworm 
but feeds on Tecoma (Bignoniaceae) (Hodges 1971). 


Macroglossinae 


Pseudosphinx tetrio (Linnaeus, 1771) 
(Frangipani hornworm) 

ID: The largest sphinx on the island; pale gray with ir- 
regular darker markings. 
DIST: United States throughout the Antilles to Ar- 
gentina. 
GUANA: 7 specimens, 1988, 1990. 
BIO: The conspicuous bright caterpillars—banded 
black and yellow with reddish brown head and orange 
legs—are frequently seen defoliating frangipani 
(Plumeria spp., Apocynaceae) on the island. 


Erinnyjis alope (Drury, 1773) 
(Papaya hornworm) 


ID: Distinguished from other sphingids on the island 
by the combination of yellow basal half of hindwing 
and alternate pairs of light gray and black dots on the 
abdomen. 

DIST: Widespread in the New World, including the 
Galapagos. 

GUANA: | specimen, 1989. 

BIO: Larvae on a variety of plants with milky sap, such as 
papaya (Carica papaya L., Caricaceae), Jatropha (Eu- 
phorbiaceae), and Alamanda (Apocynaceae). 


VOLUME 56, NUMBER 1 


Fics. 2-3. Genitalia of Catabenoides terminellus. 2, male, ventral view, aedeagus removed; 3, aedeagus, lateral view. 


Erinnyis ello (Linnaeus, 1758) 
(Cassava hornworm) 


ID: Sexually dimorphic. Male forewing dark gray with 
a blackish irregular band along the wing from near 
base to apex. Female forewing light gray with almost 
no markings. Abdomen with pairs of alternate light 
gray and black dots dorsally. 

DIST: The most common species of the genus in trop- 
ical America; also in Galapagos. 

GUANA: 5 specimens, 1984, 1990. 

BIO: Larvae varying in color from yellowish to green 
and to brownish have been serious pests of cassava 
(Manihot) in tropical America; feeds on various Eu- 
phorbiaceae. 


Erinnyis crameri (Schaus, 1898) 


ID: Forewing more brownish than those of E. ello; 
basal area reddish brown and abdomen with indistinct 
markings. 

DIST: Southern United States, through the Caribbean 
south to Brazil. 

GUANA: | specimen, 1989. 

BIO: Grayish brown larva, figured by Moss (1920: pl. 
7, figs. 3a, b), has been reared on various members of 
the Apocynaceae (Hodges 1971). 


Erinnyis domingonis (Butler, 1875) 


ID: Same size as E. obscura (see below) but forewing 


mostly dark gray. 


14 


DIST: Same as E. obscura, except for Galapagos. 
GUANA: 2 specimens, 1987. 

BIO: Unknown. 

COM: It is very likely that E. obscura and E. domingo- 
nis are only forms of the same species (Hodges 
1971:102, Kitching and Cadiou 2000: note 162). This 
could be verified by rearing. 


Erinnyis obscura (Fabricius, 1775) 


ID: Similar to E. ello, which is also dimorphic, but 
easily distinguished by its smaller size and absence of 
dots on abdomen. 

DIST: Southern United States throughout the 
Caribbean south to Brazil, including Galapagos. 
GUANA: 4 specimens, 1984, 1987. 

BIO: The pale yellowish or pale green larvae have 
been reared on various milk plants such as Philibertia 
and Cynanchum (Asclepiadaceae). 


Pachylia ficus (Linnaeus, 1758) 
(Large fig hornworm) 


ID: Large, dull brown with stout body, quite distinct 
from other sphingids on the island. Easily recognized 
by the pale, inverted trapezoidal mark on costa near 
apex. 

DIST: Widespread throughout the New World, but 
absent from Galapagos. 

GUANA: | specimen, 1987. 

BIO: Moss (1912) figured in color the several forms of 
the caterpillars, which feed on various species of Ficus 
(Moraceae). Some are green, banded yellow dorsally, 
while the others are gray brown ventrally and orange, 


banded black, dorsally. 
Callionima falcifera (Gehlen, 1943) 


ID: Recognized by the orange brown color and the 
metallic silvery mark near center of forewing. 

DIST: Southern United States to Argentina. 

GUANA: 4 specimens, 1987, 1990. 

BIO: Unknown. Other species in the genus have been 
reared on some apocynaceous plants (Hodges 1971). 
COM: Similar to, and frequently confused with, C. 
parce (Fabricius) (Kitching and Cadiou 2000: note 91). 
The specimen illustrated as C. parce in Hodges (1971: 
pl. 10, fig. 8) represents this species. 


Perigonia lusca (Fabricius, 1777) 
ID: Medium size dull brown sphinx, recognized by the 
bright orange area along the middle of hindwing. 
DIST: Southern Florida to Argentina. 
GUANA: 27 specimens, 1987, 1988, 1989, 1990. 
BIO: Bluish green larva, figured in color by Moss 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


(1912), was reared by him on coffee (Rubiaceae). 
Recorded from Gonzalagunia spicata (Lam.) Maza 
and other Rubiaceae in Puerto Rico by Torres (1992). 
COM: Kitching and Cadiou (2000: note 448) review 
names associated with P. lusca. 


Enyo lugubris (Linnaeus, 1771) 


ID: Medium size, dark brown; distinguished from 
other hawk moths on the island by the dentate borders 
of both wings. 

DIST: Southern United States, throughout the An- 
tilles, south to Uruguay and Argentina, including Gala- 
pagos. 

GUANA: 3 specimens, 1987, 1989. 

BIO: Larvae on Ampelopsis spp., Cissus spp., and Vi- 
tis spp. (Vitaceae) (Hodges 1971). 


Aellopos tantalus (Linnaeus, 1758) 


ID: Small, dark gray, diurnal, sphinx moth distin- 
guished by the conspicuous white bar across base of 
abdomen. 

DIST: New York south to Argentina. 

GUANA: One specimen captured in a Malaise trap, 
1990. 

BIO: Larvae on Ixora venulosa Benth. (Rubiaceae) 
(Biezanko et al. 1949). 


Eumorpha vitis (Linnaeus, 1758) 

(Vine sphinx) 
ID: Forewing dark green crossed with grayish bands 
and dashes, and hindwing with anal margin pink. A 
beautiful and showy species. 
DIST: United States throughout the Caribbean to Ar- 
gentina (not in Galapagos). 
GUANA: 2 specimens, 1987, 1990. 
BIO: The larvae vary in color; some are dark pink, oth- 
ers are pale green or yellow green (Moss 1912). They 
feed on grape leaves (Vitis spp.). 


Cauthetia noctuiformis (Walker, 1856) 


ID: The smallest sphingid in the New World, with a 
wing span slightly over 3 cm. Gray, with basal half of 
hindwing orange yellow. 

DIST: Caribbean. 

GUANA: 166 specimens, 1984, 1986, 1987, 1988, 
1989, 1990. 

BIO: Unknown, but larvae of C. grotei have been 
reared on Chiococca alba (L..) Hitch. (Rubiaceae) 
(Hodges 1971). 

COM: Kitching and Cadiou (2000: note 96) discuss 
the taxonomy and place the Guana population in the 
subspecies C. noctuiformis bredini Cary, 1970. 


VOLUME 56, NUMBER 1 


15 


Fics. 4-6. Genitalia of Catabenoides lazelli, new species. 4, male, ventral view, aedeagus removed; 5, aedeagus, lateral view; 6, female, 


ventral view. 


Xylophanes chiron (Drury, 1770) 


ID: Green with an oblique grayish brown irregular band 
on the forewing looking like a leaf with dead areas. 
DIST: Mexico, throughout the Antilles to Argentina. 
GUANA: 1 specimen, 1987. 

BIO: The showy caterpillar, illustrated in color by Moss 
(1920: pl. 8, figs. 6a—f), was reared by him on Palicourea, 
Psychotria, and Spermacoce (Rubiaceae). They are 
glossy green, bearing pairs of red or white red-ringed 
eye spots on the first two abdominal segments. 


Xylophanes pluto (Fabricius, 1777) 


ID: Green with irregular transverse light and dark 
bands. Recognizable by the wide orange band on the 
hindwing. 

DIST: Southern United States to Brazil. 

GUANA: 8 specimens, 1987, 1988, 1989, 1990. 


BIO: Torres (2000) described larvae from Puerto Rico, 
reared from Hamelia patens Jacq. (Rubiaceae). Gund- 
lach (1881), mentioned by Hodges (1971), also de- 
scribed the larvae, which are of two color forms: one is 
basically green, the other is black, dark violet and red. 
Gundlach (1881) recorded the hosts as Chiococca (Ru- 
biaceae) and Erythroxylum (Erythroxylaceae). 


Xylophanes tersa (Linnaeus, 1771) 


ID: Distinguished from its congeners on the island by 
the pale greenish gray forewing with several ill-defined, 
nearly parallel, longitudinal lines, running from base to 
apex; hindwing black with pale yellow marks in the vein 
interspaces parallel to the external margin. 

DIST: Ontario, across the Antilles down to Argentina, 
including Galapagos. 

GUANA: 4 specimens, 1986, 1987, 1989. 


16 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 7-9. Genitalia of Perigea gloria, new species. 7, male genitalia, ventral view, aedeagus removed; 8, aedeagus, lateral view; 9, female 


genitalia, ventral view. 


BIO: The larvae, beautifully illustrated by Moss (1912: 
pl. 14, figs. n—q), are yellowish brown with a pair of eye 
spots laterally on abdominal segments 1-7. The larvae 
feed on Psychotria berteriana DC, Borreria verticil- 
lata (L.) Meyer and Diodia sarmentosa Sw. (Rubi- 
aceae) in Puerto Rico (Torres 1992). 


Hyles lineata (Fabricius, 1775) 
(White-lined sphinx) 


ID: Resembles E. vitis, but is distinguished by its 
smaller size, shorter, clubbed antennae, and single wide 
fascia running from near base of dorsum to apex of 
forewing, crossed by whitish lines following the veins. 
DIST: This powerful flyer, almost diurnal, has reached 
all continents, as well as remote islands such as Gala- 
pagos and Hawaii. 

GUANA: I specimen, 1988. 

BIO: Polyphagous. Most commonly used plants are 


species of Portulaca (Portulacaceae), but includes oth- 
ers such as Fuchsia (Onagraceae), Boerhavia and Mi- 
rabilis (Nyctaginaceae), Xantium (Asteraceae), and 
others. Larvae are highly variable in coloration and 
somewhat in maculation. Some specimens basically 
are black with a pattern of yellow; others are mainly 
yellow with some black pattern (Hodges 1971). 


NOTODONTIDAE 


Nystalea nyseus (Cramer, 1775) 
(Fig. 15) 


ID: Narrow winged, light gray, mottled with dark 
brown and black scales; recognized by the long scales 
on the base of antennae that forms a crest on top of 
the head when resting. 

DIST: Mexico, throughout the Caribbean south to Brazil. 
GUANA: 3 specimens, 1989. 


VOLUME 56, NUMBER 1 


7 


Fics. 10,11. Male genitalia of Anateinoma affabilis. 10, ventral view, left valva and aedeagus removed; 11, aedeagus, lateral view. 


BIO: Larvae feed on various Myrtaceae especially on 
species of Psidiwm (Todd 1973:271). 

COM: This is the only notodontid present on the is- 
land; less than 30 species have been recorded from the 
Antilles (Becker in prep.). Notodontidae typically oc- 
cur in moister forests. 


NOCTUIDAE 


Heliothinae 


Heliothis subflexa (Guenée, 1852) 
(Fig. 16) 


ID: Medium sized, pale olive green; forewing crossed 
with three olive bands, edged pale basad. 


DIST: North America, throughout the Antilles, south 
to Argentina. 


GUANA: 2 males, 1989. 


BIO: Larvae on Solanum nigrum L., Physalis spp. 
(Solanaceae) (Poole et al. 1993). 


COM: Easily confused with the tobacco budworm, H. 
virescens (Fabricius), not collected but certainly oc- 
curring on the island. Male H. subflexa have white 
hindwing, while in H. virescens they are bordered 
olive-gray. More details on both species can be found 
in Poole et al. (1993). Haile et al. (1975) discuss move- 
ment of Heliothis spp. among the Virgin Islands. 


18 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics, 12-14. Genitalia of Eueana simplaria. 12, male, ventral view, left valva and aedeagus removed; 13, aedeagus, lateral view; 14, female, 


ventral view 

Noctuinae 

Anicla infecta (Ochsenheimer, 1816) 
(Fig. 17) 

ID: Gray, with forewing reddish brown along external 
margin; hindwing hyaline. Recognized by blackish an- 
terior border (patagia) of thorax. 
DIST: Argentina through Central United States, in- 
cluding Galapagos and Bermuda. 
GUANA: 6 specimens, 1986, 1989, 1990. 
BIO: General feeder, cut worm. The larvae reach 
nearly 3 cm when fully grown. They are variable in 
color from gray to yellowish ferrugineous, olivaceous 
yellow and bright green, to a sordid brown, usually 


flecked with black. 


COM: At least three other cut worms should be present 
in the island: Agrotis ipsilon (Hufnagel), A. subterranea 
(Fabricius) and Peridroma saucia (Hiibner). 


Hadeninae 


Leucania humidicola Guenée, 1852 
(Fig. 19) 


ID: Medium sized, pale moth. Distinguished from 
other noctuids on the island by its forewing pattern: a 
long dark dash delimited above by a white line, run- 
ning from base to middle. 

DIST: Antilles to Brazil and probably Galapagos; the 
limits of the distribution of the species remain to be 
determined (Adams 2001). 


VOLUME 56, NUMBER 1 19 


Fics. 15-61. Natural size (1:1). Notodontidae (15) and Noctuidae (16-61) (species from Guana, unless stated otherwise). 15, Nystalea ny- 
seus, male; 16, Heliothis subflexa, male (USA); 17, Anicla infecta, male; 18, Leucania dorsalis, female (Cuba); 19, L. humidicola, male; 20, 
Neogalea sunia, male; 21, Catabenoides lazelli, holotype male; 22, C. terminellus, female; 23, Spodoptera albulum, male; 24, S. frugiperda, ie 
walle (Puerto Rico); 25, S. PS ee male (Brazil); 26, S. latifascia, male; 27, S. latifascia, female (Cuba); 28, S. pulchella, male; 29, 
dolichos, male (Mexico); 30, Magusa orbifera, female; 31, Condica albigera, rele 32, C. albigera, female; 33, C. mobilis, male (Brazil); C. su- 
tor, male (Brazil); 35, ee gloria, holotype male; 36, Elaphria agrotina, male; 37, E. aedgalloria, female; 38, E. nucicolora, male (Cuba); 39, 
Micrathetis triplex, female; 40, M. triplex, male; 41, Bagisara repanda, female; 42, Amyna axis, female; 43, Ponometia exigua, male; 44, P. ex- 
igua, female; 45, P. exigua, female (Mexico); 46, Cydosia nobilitella, male (Cuba); 47, Caularis undulans, male; 48, Motya Aaa alis, male: 49, 
Collomena filifera, male; 50, C. filifera, female (Cuba): 51, Paectes obrotunda, male; 52, P. obrotunda, female; 53, Pseudoplusia includens, fe- 
male; 54, Ptichodis immunis, male (Cuba): 55, P. immunis, female; 56, Mocis antillesia, male; 57, M. antillesia, female; 58, M. latipes, male; 59, 
M. repanda, male (Cuba); 60, M. repanda, female (Puerto Rico); 61, Ophisma tropicalis, male. 


GUANA: 2 specimens, 1989. 

BIO: Hayes (1975) [as L. solita, see below] gives 
Sporobolus virginicus (L.) Kunth. (Poaceae) as food- 
plant. 

COM: The Guana specimens were identified as hu- 
midicola by Morton S. Adams, who has subsequently 
published a revision of the group in the Caribbean 
(Adams 2001) although our specimens are not men- 
tioned in his paper. The name hwmidicola (type-locality: 
FRENCH GUIANA) has been wrongly applied to a dif- 
ferent species by most authors (see dorsalis below), fol- 
lowing the misidentification by Hampson (1905). A 
specimen collected by the first author in COSTA RICA: 
Guanacaste, E] Coco (VOB 33636), identical to those 
from Guana, matches the type of solita (type-locality: 
HONDURAS) in BMNH, and the specimen figured by 
Hayes (1975), from Galapagos. Therefore L. solita 
Walker 1856, new synonym, is a junior synonym of hu- 
midicola, not of multilinea Walker (sensu Hampson 
1905, Poole 1989). We regard multilinea Walker 1856 as 
a valid species because we believe that Hayes (1975), 
who had the types of both multilinea and solita at hand, 
had good reasons to treat the latter as a valid species. 


Leucania dorsalis Walker, 1856 
(Fig. 18) 


ID: Easily confused with L. humidicola; dash along 
middle of forewing not as conspicuous. 
DIST: Antilles, northern South America, Central 
America, and southern Florida (Adams 2001). 
GUANA: | specimen, 1988. 
BIO: Unknown; presumably grasses as for its close rel- 
atives L. infatuans Franclemont and L. extenuata 
Guenée. 
COM: This species belongs to a complex previously 
treated as humidicola, following a misidentification by 
Hampson (1905). See Adams (2001:199) for further 
discussion of the species complex. 
Amphipyrinae 
Neogalea sunia (Guenée, 1852) 
(Fig. 20) 
ID: Medium sized, inconspicuous gray moth resem- 
bling Spodoptera albulum, but darker. Usually distin- 
guished from S. albulum by the dark marked veins 
and, for males, by the abdomen thickly clothed with 
long scales. 
DIST: Florida to Argentina, including Galapagos. 
GUANA: 6 specimens, 1987, 1989, 1990. 
BIO: Larvae on Lantana (Verbenaceae) (Comstock 
and Dammers 1935). Introduced to Hawaii and Aus- 


tralasia to control Lantana (Riotte 1991, Holloway 
1996:148). 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


COM: Immature stages described by Comstock and 
Dammers (1935, as Catabene esula). Although this 
genus was placed in Cuculliinae by Poole (1989), we 
place it in Amphipyrinae following the comments by 
Todd (1972b) and Poole (appendix to this paper) plac- 
ing it with Catabena and Catabenoides, which Poole 
(1989) placed as Amphipyrinae. The proper placement 
of many genera formerly associated with Amphipyri- 
nae and Cuculliinae requires review. 


Catabenoides terminellus (Grote, 1883), new 
combination 
(Figs. 2, 3, 22) 


ID: This and the following species are closely related, 
almost impossible to distinguish with external charac- 
ters (see C. lazelli below). They resemble small S. al- 
bulum but are distinguished from it by the gray, irreg- 
ular dot on tornus of forewing. Females usually have a 
black line along middle, covering the length of the 
forewing. 

DIST: Southern USA, Antilles. 

GUANA: 4 specimens, 1984. 

BIO: Unknown. 

COM: In order to place this and the following species 
correctly, we include a description of the new genus 
Catabenoides by Robert Poole as an appendix to this 
paper. 


Catabenoides lazelli Becker and Miller, new 
species 
(Figs. 4-6, 21) 


Description. Light gray, 2.2-2.6 cm. This and the 
former are very closely related species, almost impos- 
sible to be distinguished from each other on external 
characters (see terminellus above). They resemble a 
small S$. albulum but easily recognized from it by the 
gray, irregular dot on tornus of forewing. Females usu- 
ally have a black line along middle, covering the whole 
extension of forewing. The only reliable external fea- 
ture that distinguishes lazelli from terminellus is the 
color of patagia. In terminellus there is a transverse 
line of blackish scales, dividing the patagia along the 
middle, while in lazelli the line is ochreous. The geni- 
talia are also distinct. In terminellus the distal 
processes of the sacculus are simple, nearly straight 
rods (Fig. 2), while those in lazelli are complex, 
branched (Fig. 4). 

DIST: Guana, Anegada, St. Croix. 

GUANA: 35 specimens, 1984, 1985, 1989, 1990. 
MATERIAL EXAMINED: Holotype male: BVI: 
Guana Id., 1-14.vii.1984 (S. E. & PB M. Miller) 
(USNM). Paratypes: 13 males, 12 females: Same data 
as holotype (USNM, BMNH, BPBM, MCZ, VOB); 1 


VOLUME 56, NUMBER | 


female: Same locality and collector, 5—23.vii.1985 
(USNM); 2 males, 1 female: Same locality, x.1989 (V. 
O. Becker, 70710) (VOB); 2 males, 3 females: Same 
locality, 24.x-5.xi.1990 (S. E. Miller & T. M. Kuklen- 
ski) (BPBM). ANEGADA: 2 males, 1 female: 
17-19.vii.1985 (S. E. & BP M. Miller) (USNM). ST. 
CROIX: 2 males, 2 females, Kingshill, x, xi.1944, v, 
vi.1945 (H. A. Beatty) (CU); 1 female, same locality, 
6-16.vii.1967 (E. L. Todd) (USNM); 1 female, Chris- 
tiansted, 19. xi.1941 (H. A. Beatty) (USNM); 1 male, 2 
females, Mt. Eagle, 6-16.vii.1967 (E. L. Todd) 
(USNM); 4 males, 1 female, 1 mi W Airport, 
6—-16.vii.1967 (E. L. Todd) (USNM); 2 males, 1 fe- 
male, Orangegrove, W End, 6-16.vii.1967 (E. L. 
Todd) (USNM): 1 male, Blue Mtn., 6—16.vii.1967 (E. 
L. Todd) (USNM); 2 males, Rust Up Twist, 
6-16.vii.1967 (E. L. Todd) (USNM): 1 male, 1 mi N 
Great Pond, 6—16.vii.1967 (E. L. Todd) (USNM). 
BIO: Unknown. 

COM: This species belongs to a complex formerly con- 
sidered the single species, C. vitrinus (Walker), a 
species not found in the Lesser Antilles. The genitalia 
of C. lazelli (Fig. 4, 5) are very similar, but show con- 
sistent differences, the most evident is the vesica 
armed with a single, strong cornutus, whereas in vitri- 
nus the vesica bears a series of smaller cornuti. The 
complex will be treated in a forthcoming revision 
(Becker in prep.). This species is dedicated to our 
friend Dr. James “Skip” Lazell, who gave us the op- 
portunity to study this interesting fauna. 


Spodoptera albulum (Walker, 1857) 
(Fig. 23) 


ID: Plain, pale gray, medium sized species, readily dis- 
tinguished by the presence, on the forewing, of a very 
fine black line running along the middle from base to 
one-fourth. Hindwing almost totally translucent whitish. 
DIST: United States, throughout the Antilles, south to 
Argentina, but not including Galapagos and Bermuda. 
GUANA: 5 specimens, 1989, 1990. 

BIO: Larvae on Amaranthus sp. (Amaranthaceae) 
(Kimball 1965) and cotton (Bruner et al. 1975). 
Recorded from many crops in Puerto Rico by Arm- 
strong (1994a). 

COM: “This is the species previously identified as 
‘Spodoptera sunia Guenée’. The real Xylopmyges Su- 
nia Guenée 1852 is actually the species [formerly] 
known as Neogalea esula Druce” (Poole 1989) (see N. 
sunia above). Spodoptera albulum is easily confused 
with S. eridania, which has not been collected on 
Guana, but is likely to occur on the island. Spodoptera 
eridania is dusted brownish, and lacks the forewing 
line mentioned above. Todd and Poole (1980) give an 


2 


illustrated key to the New World species of Spo- 
doptera and distributions of Spodoptera species in the 
Caribbean are reviewed in Cock (1985:92). 


Spodoptera frugiperda (J. E. Smith, 1797) 
(Fall armyworm) 
(Figs. 24, 25) 


ID: Medium sized, sexually dimorphic, gray species. 
Males have an oblique whitish dash from middle of 
costa across the cell. Females have indistinct pattern, 
looking almost plain gray. 

DIST: Widespread in New World, including Galapa- 
gos and Bermuda. 

GUANA: 8 specimens, 1986, 1990. 

BIO: Polyphagous on herbaceous plants and regarded 
as a serious pest of maize and other crops (Andrews 
1980). In Puerto Rico, it has been recorded as a pest of 
various crops (Armstrong 1994b) and Eucalyptus 
seedlings (Myrtaceae) (Torres 1994). 


Spodoptera latifascia (Walker, 1856) 
(Figs. 26, 27) 


ID: Medium sized, sexually dimorphic species. Male 
forewing with a diffuse pattern of reddish brown and 
gray on a whitish gray background. Female forewing 
darker, easily confused with S. dolichos and bearing an 
oblique elongate whitish mark from middle costa to 
end of cell, followed by three short whitish lines along 
veins. 

DIST: Gulf States of the United States, throughout the 
Antilles, south to Costa Rica. The population from 
Costa Rica south to Argentina, previously included un- 
der S. latifascia, belongs to S. cosmioides (Walker), a 
closely related but distinct species (Silvain & Lalanne- 
Cassou 1997, M. Pogue pers. com.). 

GUANA: | specimen, 1989. 

BIO: Polyphagous on herbaceous plants, sometimes 
becoming a pest of vegetables and nursery seedlings. 


Spodoptera pulchella (Herrich-Schiffer, 1868) 
(Fig. 28) 


ID: Wing pattern similar in both sexes; easily confused 
with the females of the former. It can be separated 
from similar species by the curved whitish line along 
dorsum, below the anal vein, from basal fourth to just 
before tornus. 

DIST: Florida, Greater Antilles. 

GUANA: | specimen, 1989. 

BIO: Unknown. 

COM: This seems to be the first record of this species 
to the Puerto Rican Bank. It has either been over- 
looked because of rarity, or because it was mistaken for 
the similar S. latifascia, a more common species. 


bo 
ins) 


Spodoptera dolichos (Fabricius, 1794) 
(Fig. 29) 

ID: About the same size as S. latifascia; both sexes show- 
ing similar pattern to that of female S. latifascia. Easily 
distinguished from the previous two species by the two 
conspicuous, parallel, dark gray bands along thorax. 
DIST: Sympatric with S. latifascia, including in Gala- 
pagos. 
GUANA: 1 specimen, 1989. 
BIO: Larvae on a wide variety of plants, both crops 
and weeds (Ferguson et al. 1991). 


Magusa orbifera (Walker, 1857) 
(Fig. 30) 


ID: An extremely polymorphic, medium sized (3-4 
cm), gray to brown species. In the Guana population, 
some males have a wide pale area along dorsum of 
forewing, others have a very complex and contrasting 
maculation, while the females tend to be less marked 
and more brownish. One constant feature is the con- 
spicuous round pale dot near the apex of forewing and 
the very broad dark fuscous hindwing. Anateinoma af- 
fabilis and E. agrotina also have the pale mark at end 
of apex of forewing but are at most half the size of M. 
orbifera. 

DIST: Widespread throughout the New World, from 
Canada to Argentina (not reported from Galapagos, 
but M. erema Hayes (1975) may be a local variety of 
this species). 

GUANA: 57 specimens, 1989, 1990. 

BIO: Larvae on various legumes, including Karwin- 
skia and Condalia (Fabaceae) (Kimball 1965). 


Condica albigera (Guenée, 1852) 

(Figs. 31, 32) 
ID: Medium sized (2.5-3 cm wing span), dark fuscous; 
forewing with an irregular small white dot at end of 
cell, followed by a paler, almost straight transverse line. 
DIST: Mexico, throughout Antilles, south to Paraguay. 
GUANA: 3 specimens, 1989, 1990 
BIO: Unknown. 
COM: Easily confused with C. circuita (Guenée), not 
collected but likely to occur on the island. In C. cir- 
cuita the white dot on forewing is round and has a 
white lunule just under it. 


Condica mobilis (Walker, [1857]) 
(Fig. 33) 


ID: About same size as C. albigera, but more reddish 
brown and orange; white dot on cell usually larger 
than in C. albigera. 
DIST: Southern United States, throughout Antilles, 
south to Argentina. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


GUANA: | specimen, 1989. 

BIO: Unknown. 

COM: Commonly referred to in the literature as 
Perigea apameoides Guenée, which is a synonym of C. 
sutor (Guenée) (Hayes 1975). 


Condica sutor (Guenée, 1852) 
(Fig. 34) 


ID: Same size as C. albigera and C. mobilis; fuscous 
with forewing showing little contrasting pattern. Dis- 
tinguished from albigera and mobilis by the absence of 
the whitish mark on cell. 

DIST: Southern United States, throughout the An- 
tilles, south to Argentina, including Galapagos. 
GUANA: | specimen, 1990. 

BIO: Larvae on several species of herbaceous plants 
including Wedellia, Tagetis (Asteraceae), and celery 
(Kimball 1965). 


Perigea gloria Becker and Miller, new species 
(Figs. 7-9, 35) 


Description. Medium sized (3 cm wing span), pale 
moth; forewing shaded dark fuscous, with a series of 
small dark marks along costa and small black dots 
along termen, in the spaces between veins. Similar to 
C. sutor but with more contrasting pattern, and read- 
ily separated by the series of black dots along termen. 
DIST: Guana, Tortola. 

GUANA: 2 specimens, 1989. 

MATERIAL EXAMINED: Holotype male: BVI: 
Guana Id., x.1989 (V. O. Becker, 70722) (USNM): 
Paratypes, 1 female, same data as holotype (VOB); 1 
male Tortola, Mt. Sage, 460m, 13—-15.vii.1987 (V. O. 
Becker & S. E. Miller, 66865) (VOB). 

BIO: Unknown. 

COM: Very similar in appearance to P. berinda 
(Druce), a species from the Greater Antilles and Cen- 
tral America, but with genitalia (Figs. 7-9) very differ- 
ent from those of berinda, being very similar to those 
of P. glaucoptera (Guenée). This species is dedicated 
to Ms. Gloria Jarecki, for her and her family’s support 
of The Conservation Agency's biodiversity research on 
Guana Island over the years. 


Elaphria agrotina (Guenée, 1852) 
(Fig. 36) 


ID: Small (2—-2.5 cm wing span); forewing dark fuscous 
with a paler area along costa and a conspicuous pale 
dash near apex. Similar to A. affabilis (see below), but 
larger and with hindwing bordered whitish. 

DIST: Florida, throughout the Antilles, south to Ar- 
gentina. 


VOLUME 56, NUMBER 1 


GUANA: 5 specimens, 1987, 1989, 1990. 
BIO: Larvae on cotton and beans (Phaseolus) 
(Fabaceae) (Silva et al. 1968). 


Elaphria nucicolora (Guenée, 1852) 
(Figs. 37, 38) 


ID: Same size as E. agrotina; forewing dark fuscous, 
with a broad, ill-defined, darker triangular mark with 
base on middle of dorsum and vertex at end of cell. 
Hindwing whitish. 

DIST: Throughout New World tropics including 
Bermuda. Immigrant to Hawaii. 

GUANA: | specimen, 1989. 

BIO: Larvae on various herbaceous plants (Ferguson 
et al. 1991). 


Anateinoma affabilis Méschler, 1890 
(Figs. 10, 11, 151, 152) 


ID: Small (15 mm wing span); forewing reddish brown 
with transverse sinuate lines alternating pale and dark, 
and with a conspicuous whitish dash on apex. Similar 
to E. agrotina (see above) but smaller, and forewing 
lacking pale area along costa. Magusa orbifera also has 
a pale mark on apex, but is almost three times the size 
of affabilis. 

DIST: Puerto Rico and Virgin Islands. 

GUANA: 22 specimens, 1987, 1989, 1990. 

BIO: Unknown. 

COM: Despite the accurate color illustration pre- 
sented by Méschler (1890), who described this species 
from Puerto Rico, Hampson (1910) treated A. affabilis 
as an unrecognized taxon in the Erastriinae |=Aconti- 
inae], where it has remained. One of the reasons for 
this situation is because no material except for the 
types, which are supposed to be in MNHU, Berlin, has 
been available to subsequent authors working on the 
New World noctuid fauna. No material of this species 
was found in the BMNH or USNM,; for this reason 
vouchers from the series studied here have been de- 
posited there. 

This species does not belong in Acontiinae, but is 
related to some species currently placed in Elaphria 
Hiibner. However, at present we prefer not to syn- 
onymize Anateinoma under Elaphria as the group 
needs revision. The male genitalia is illustrated in Figs. 
10, 11. 


Micrathetis triplex (Walker, 1857) 
(Figs. 39, 40) 


ID: Small (1.5-2.2 em wing span), slightly dimorphic, 
variable in color. Males have pale forewing speckled 
with darker small dots, termen dark brown, and a con- 
spicuous dark brown dot at end of cell. Females are 


darker than males. Hindwing whitish, slightly bor- 
dered with dark gray. Easily recognized by the dot at 
the end of the cell and by the two rows of small black- 
ish dots forming two arches, almost parallel to each 
other, from costa to dorsum. 
DIST: Southern United States through South America. 
GUANA: 14 specimens, 1987, 1989, 1990. 
BIO: Unknown. 
Agaristinae 
Caularis undulans Walker, [1858] 
(Fig. 47) 
ID: Undoubtedly the most attractive noctuid on the is- 
land. Forewings white bordered and marked olive; 
hindwing golden yellow bordered reddish-brown with 
a lunular blackish mark on tornus. Male genitalia illus- 
trated by Kiriakoff (1976). 
DIST: Hispaniola, Jamaica, Puerto Rican Bank (Kiri- 
akoff 1976). 
GUANA: 9 specimens, 1989. 
BIO: Unknown. 


Bagisarinae 


Bagisara repanda (Fabricius, 1793) 
(Fig. 41) 


ID: Small, 2-2.5 cm wing span, pale yellow, dusted 
gray. Easily identified by the three pale lines crossing 
the forewing, more or less equidistant, parallel to each 
other, and bent basad near costa. 

DIST: Widespread from Southeast United States to 
Paraguay, including Galapagos (Hayes 1975, Ferguson 
1997). 

GUANA: 4 specimens, 1987, 1989. 

BIO: Larvae on Sida glomerata Cav. (Malvaceae) 
(Hayes 1975). 

COM: Very common in disturbed areas where malva- 
ceous weeds often occur. 


Acontiinae 


Amyna axis (Guenée, 1852) 
(Fig. 42) 

ID: Small, fuscous species, easily confused with some 
small Condica species. Males are distinguished from 
Condica by the presence of a round, semitranslucent 
area near base of forewing. 
DIST: Pantropical, including Tahiti and Hawaii. 
GUANA: 2 specimens, 1989. 
BIO: Larvae on Chenopodium (Chenopodiaceae), 
Cardiospermum (Sapindaceae), Parasponia (UI- 
maceae), and Amaranthus (Amaranthaceae) (Fergu- 
son 1991). 
COM: The large distribution and obscure pattern have 
contributed to long synonymy: it has been described 


94 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 62-111. Natural size (1:1). Noctuidae (specimens from Guana, unless stated otherwise). 62, Azeta versicolor, male; 63, A. versicolor, 
female; 64, 65, Metallata absumens, males; 66, M. absumens, male; 67, Plusiodonta thomae, male; 68, Syllectra erycata, male (Cuba); 69, Lito- 
prosopus puncticosta, male; 70, Diphthera festiva, male; 71, Gonodonta bidens, male (Puerto Rico); 72, Melipotis acontioides, female; 73, M. 
fasciolaris, male; 74, M. fasciolaris, female; 75, M. contorta, male; 76, M. famelica, male; 77, M. ochrodes, male; 78, M. ochrodes, female 
(Puerto Rico); 79, M. januaris, male (Cuba); 80, M. januaris, female (Cuba); 81, Epidromia lienaris, male (Puerto Rico); 82, Ephyrodes cacata, 
male (Cuba); 83, E. cacata, female; 84, Concana mundissima, female; 85, Massala asema, male; 86, Manbuta pyraliformis, male (Cuba); 87, 
Lesmone hinna, male (Cuba); 88, L. hinna, female (Cuba); 89, L. hinna, male; 90, L. formularis, male; 91, L. formularis, female (Cuba); 92, 
Baniana relapsa, male; 93, B. relapsa, female; 94, Eulepidotis modestula, male (Cuba); 95, E. addens, female; 96, Toxonprucha diffundens, 
male (Mexico); 97, Kakopoda progenies, male; 98, Parachabora abydas, male; 99, Cecharismena abarusalis, male; 100, C. cara, male; 101, 
Glympis eubolialis, male; 102, Drepanopalpia lunifera, male (Cuba); 103, D. lunifera, female; 104, Lascoria orneodalis, female; 105, L. or- 
neodalis, male (Cuba); 106, Bleptina caradrinalis, male; 107, B. caradrinalis, female; 108, B. hydrillalis, male; 109, B. menalcasalis, female; 
110, B. menalcasalis, male; 111, Hypena lividalis, female. 


VOLUME 56, NUMBER 1 25 


‘ 

é {> * Ss 
ee Ly 

a a ™ 
Se ee 


FAY 


ee —< S oon 
i Coys shi eo aN 


Fics. 112-150. Natural size (1:1). Arctiidae (112-124), Geometridae (125-146), Cossidae (147-149) and Hyblaeidae (150) (specimens from 
Guana, unless stated otherwise). 112, Hypercompe simplex, male (Puerto Rico); 113, Composia credula, male; 114, H. simplex, female; 115, 
Calidota strigosa, male; 116, Eupseudosoma involutum, male (Puerto Rico); 117, Utetheisa ornatrix, male; 118, U. pulchella, female (Brazil); 
119, Empyreuma pugione, male; 120, Horama panthalon, male; 121, H. pretus, male; 122, Cosmosoma achemon, male (St. Thomas); 123, Eu- 
nomia colombina, male; 124, Nyridela chalciope, female (Cuba); 125, Pero rectisectaria, male; 126, P. rectisectaria female; 127, Oxydia vesu- 
lia, male; 128, Erastria decrepitaria, male (Cuba); 129, E. decrepitaria, female; 130, Sphacelodes fusilineatus, male; 131, S. fusilineatus, fe- 
male; 132, Macaria paleolata, male; 133, Patalene ephyrata, male; 134, Almodes terraria, male (Bahamas); 135, Semaeopus malefidarius, male; 
136, Leptostales noctuata, male; 137, L. noctuata, female; 138, Obila praecurraria, female (Tortola); 139, Pterocypha defensata, male; 140, P. 
defensata, female; 141, Eueana simplaria male; 142, E. simplaria female; 143, Phrudocentra centrifugarium, male; 144, P. centrifugarium, fe- 
male (Cuba); 145, 146, P. centrifugarium, females; 147-149, Psychonoctua personalis, males; 150, Hyblaea puera, male (Cuba). 


18 times (Poole 1989). Frequently referred to in the 
literature as A. octo (Guenée), a synonym based on the 
priority of names established by Nielsen et al. (1996: 
note 690). 


Ponometia exigua (Fabricius, 1793) 
(Figs. 43-45) 


ID: Small, variable, sexually dimorphic species; males 
pale yellow with forewing crossed with diffuse, sinuate 
olivaceous bands. Female forewing dark fuscous with a 
wide, contrasting, pale fascia along costa. In some fe- 
males this pattern is less contrasting. 

DIST: Southern United States, throughout the An- 
tilles, south to Brazil, including Galapagos. 

GUANA: 9 specimens, 1989, 1990. 

BIO: Larvae on Waltheria ovata Cav. (Sterculiaceae) 
(Hayes 1975). 

COM: Commonly referred to in the literature by its 
junior synonym P. indubitans (Walker). 


Cydosia nobilitella (Cramer, 1779) 
(Fig. 46) 


ID: Small showy moth; one of the most attractive noc- 
tuids on the island. Forewing with reticulated pattern 
with white areas enclosed by dark bluish metallic gray 
and red lines. Hindwing semitranslucent white in 
males, dark gray in females. Pattern resembles some 
species of Atteva (Yponomeutidae) and specimens are 
often found mixed in collections. 

DIST: Southern United States, throughout Antilles, 
south to Argentina. 

GUANA: | specimen, 1984. 

BIO: Hampson (1910) mentioned “?Spigelia an- 
thelmia L.” (Loganiaceae) as hostplant, following 
Cockerell (1897). No species of this plant family 
known from the island (G. Proctor pers. com.), al- 
though the species occurs on other Virgin Islands 
(Acevedo-Rodriguez 1996). Cockerell (1897) and Dyar 
(1897) described the larvae. 


Tripudia quadrifera (Zeller, 1874) 
(Figs. 153, 154) 


ID: The smallest noctuid on the island (0.7—1.2 cm 
wing span); resembling Olethreutinae (Tortricidae) 
and C. metaspilaris (see below). Dark gray; forewing 
with conspicuous quadrate mark on middle of dorsum. 
DIST: Southern United States, throughout the An- 
tilles, south to Brazil. 

GUANA: 22 specimens, 1989, 1990. 

BIO: Unknown. 


Tripudia balteata Smith, 1900 
(Fig. 155) 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


ID: Small, on average slightly larger than T. 
quadrifera. Dark gray. Easily identified by the broad, 
oblique, yellowish band on forewing. 

DIST: Southern United States, Antilles, south to 
Brazil. 

GUANA: 16 specimens, 1987, 1989, 1990. 

BIO: Unknown. 


Ommatochila mundula (Zeller, 1872) 
(Fig. 156) 


ID: Small, 1.5—2 cm wing span, dark gray, resembling 
some Olethreutinae species (Tortricidae). Forewing 
divided across the middle by a pale, almost straight 
line, the basal half much darker than outer half. 
DIST: Southern United States, Antilles, south to Ar- 
gentina. 

GUANA: 14 specimens, 1986, 1989, 1990. 

BIO: Unknown. 


Cobubatha metaspilaris Walker, 1863 
(Fig. 157) 


ID: Small, 1.5 cm wing span, gray; similar to, but 
larger than T. quadrifera. In the latter the mark on 
dorsum is quadrate whereas in metaspilaris it is trape- 
zoidal. 

DIST: Antilles. 

GUANA: 3 specimens, 1990. 

BIO: Unknown. 


Eumicremma minima (Guenée, 1852) 
(Fig. 158) 


ID: Very small (1.2-1.5 cm wing span); forewing pale, 
crossed with olive and dark olive waiving bands, and 
with some very small, black dots along termen, the 
most conspicuous the one near apex and the other 
near tornus. In resting posture it looks like some 
species of Cochylini (Tortricidae). 

DIST: Southern United States, Antilles, south to Ar- 
gentina. 

GUANA: 3 specimens, 1987. 

BIO: Larvae on Gnaphalium (Asteraceae). 


Eublemma rectum (Guenée, 1852) 
(Fig. 161) 


ID: Small, 1.5 cm wing span; pale yellow; forewing 
clouded with red brown with oblique pale fascia from 
middle of dorsum to near apex. 

DIST: Southern United States, throughout the An- 
tilles, south to Argentina, including Galapagos and 
Bermuda. 

GUANA: 3 specimens, 1989. 

BIO: Larvae on Ipomoea and Convolvulus (Convolvu- 


VOLUME 56, NUMBER | 


12) 
~ 


Figs. 151-188. Twice natural size (2:1). Noctuidae (151-168), Arctiidae (169-171) and Geometridae (172-188) (specimens from Guana, un- 
less stated otherwise). 151, 152, Anateinoma affabilis, males; 153, Tripudia quadrifera, female (Mexico); 154, T. quadrifera, female; 155, T. 
balteata, male; 156, Ommatochila mundula, female; 157, Cobubatha metaspilaris, male; 158, Eumicremma minima, male (Cuba); 159, 
Spragueia margana, male, (Brazil); 160, S. margana, female (Brazil); 161, Eublemma rectum, male; 162, Thioptera aurifere, male (Brazil); 163, 
Characoma nilotica, female; 164, 165, C. nilotica, females (Mexico); 166, Hypena minualis, female; 167, Bleptina araealis, male; 168, B. 
araealis, female; 169, Afrida charientisma, male; 170, Progona pallida, male; 171, Lomuna nigripuncta, female; 172, “Idaea” monata, male, 
173, Idaea monata, female; 174, Idaea eupitheciata, female, 175, I. eupitheciata, male; 176, I. minuta, male; 177, I. minuta, female; 178, 
Scopula laresaria, female; 179, 180, Idaea probably fernaria, females; 181, Leptostales phorcaria, male; 182, Cyclomia mopsaria, male; 183, 
C. mopsaria, female; 184, Leptostales oblinataria, female; 185, Acratodes suavata, male; 186, Chloropteryx paularia, male; 187, Synchlora 


frondaria, male; 188, S. cupedinaria, male. 


laceae) (Forbes 1954) [as E. obliqualis (Fabricius), a 
homonym]. 


Spragueia margana (Fabricius, 1794) 
(Figs. 159, 160) 


ID: The smallest of the two Spragueia species on the 
island; dimorphic. Similar to S. perstructana (see be- 
low). Males easily distinguished from the latter by the 
absence of orange, by the pale costa, and by the oliva- 
ceous shades and marks on forewing; females by the 
olivaceous thorax, which is edged pale yellow in S. per- 
structana. 


DIST: Southern United States, throughout the New 
World tropics, including Galapagos. 

GUANA: 4 specimens, 1987. 

BIO: Larvae on Abutilon and Sida (Malvaceae) (Hayes 
1975). 


Spragueia perstructana (Walker, 1865) 


ID: Similar to, but slightly larger than S. margana (see 
above); also dimorphic. Illustrated in color in Kimball 
(1965: PI. IV; figs. 31, 37). 

DIST: Southern United States, Antilles, south to Costa 
Rica. 


GUANA: 1| specimen, 1989. 
BIO: Unknown. 


Thioptera aurifere (Walker, [1858]) 
(Fig. 162) 


ID: Small, 1.5-1.8 cm wing span; yellow; forewing 
usually with two very small black dots, an ill defined 
reddish line beyond the cell from costa to dorsum, and 
termen edged with gray. 

DIST: Southern United States, throughout Antilles, 
south to Brazil. 

GUANA: 4 specimens, 1989, 1990. 

BIO: Unknown, however Kimball (1965) gives Digi- 
taria ischaemum [Syntherisma impomoea] (Poaceae) 
as the food plant for T. nigrofimbria, a closely related 
species. 


Sarrothripinae 


Characoma nilotica (Rogenhofer, 1882) 
(Figs. 163-165) 


ID: Small, 1-1.2 cm wing span, highly variable, gray 
species. Rests flat, looking like some tortricids. 

DIST: Described from Egypt, hence its name; now 
Pantropical, including Galapagos, Bermuda, and the 
Pacific Islands. 

GUANA: 25 specimens, 1985, 1986, 1990. 

BIO: Larvae on white mangrove, Laguncularia race- 
mosa (L.) C.F. Gaertn. (Combe (Hayes 1975); 
willow, almond, azalea, and “black olive” (Ferguson 1991). 


Collomena filifera (Walker, 1857) 
(Figs. 49, 50) 


ID: Medium sized gray species, similar to female S. 
frugiperda. Distinguished by the whitish diffused band 
across the subter saa area of forewing and by the en- 
tirely whitish, semitranslucent hindwing (narrowly 
bordered gray in females). 

DIST: Florida, throughout the Antilles, south to 
Brazil. 

GUANA: 4 specimens, 1989, 1990. 

BIO: Unknown. 


Motya abseuzalis Walker, 1859 
(Fig. 48) 

ID: Slightly smaller than C. filifera; whitish gray. Easily 
recognized by the two, almost parallel, rows of small, 
black dots along termen, with the one near tornus con- 
spicuously larger. The male abdomen has two paired 
black dots dorsally, near apex. Hindwing semitranslu- 
cent white, bordered gray. 

DIST: Florida, throughout the Antilles, south to 
Brazil. 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


GUANA: | specimen, 1989. 
BIO: Unknown. 


Euteliinae 


Paectes obrotunda (Guenée, 1852) 
(Figs. 51, 52) 


ID: Medium sized, gray, irrorated brown; males have 
basal half of antennae strongly pectinate, and long slim 
abdomen; females have filiform antennae and short 
stout abdomen. Forewing with a conspicuous pale 
lunular mark near base, delimited externally by a nar- 
row, double line. 

DIST: Southern United States, throughout Antilles, 
south to Paraguay. 

GUANA: 64 specimens, 1984, 1987, 1988, 1989, 1990. 
BIO: Unknown, however, its larvae should be searched 
for on Bursera simaruba (L.) Sarg. (Simarubaceae), as 
a related species, P. arcigera (Guenée), was reared on 
B. graveolens (Kunth) Triana & Planch. in Galapagos 
(Hayes 1975). 


Plusiinae 


Pseudoplusia includens (Walker, [1858]) 
(Fig. 53) 


ID: Medium sized, grayish brown with bronze luster. 
Recognized by the small silver markings near center of 
forewing. 

DIST: United States to northern Chile and Argentina, 
including Galapagos and Bermuda (atone & 
Poole 1991: 50). 

GUANA: 2 specimens, 1989. 

BIO: Polyphagous; Ferguson et al. (1991) lists plants 
belonging to 14 families as foodplants. Can be a minor 
pest of beans, soy beans, and other leguminous crops. 
COM: Generally referred to in the literature as P. 00 
(Cramer), a homonym. Other species belonging to this 
subfamily, such as Trichoplusia ni (Hiibner) and Argy- 
rogramma verruca (Fabricius), are likely to be col- 
lected on the island in the future. These also bear sil- 
ver marks on the forewing. 


Catocalinae 


Ptichodis immunis (Guenée, 1852) 
(Figs. 54, 55) 


ID: Medium sized, 2.5-3 cm wing span, pale species. 
Forewing crossed with ill defined olivaceous lines, two 
of them highly contrasting: the antemedial and the 
postmedial, both bordered internally with lemon yel- 
low. 

DIST: Mexico, throughout the Antilles to Brazil. 
GUANA: 6 specimens, 1987, 1989, 1990. 

BIO: Unknown. 


VOLUME 56, NUMBER | 


Mocis latipes (Guenée, 1852) 
(Fig. 58) 


ID: Medium sized, 3.54 cm wing span, broad winged, 
dark species. Highly variable in color and pattern. 
Ground color varies from pale brownish through fus- 
cous to reddish brown. Females tend to have pattern 
less contrasting than males, and the paler forms could 
easily be confused with the darker forms of female M. 
disseverans, a Neotropical species recorded from the 
Greater Antilles. Smaller than M. repanda (see below). 
The males of the species belonging to this genus can 
be easily distinguished from other noctuids by the 
thickly hairy hind legs. 

DIST: Southern United States, throughout the An- 
tilles, south to Argentina, including Galapagos and 
Bermuda. 

GUANA: | specimen, 1989. 

BIO: Larvae on several species of grasses, sometimes a 
pest of grazing land. 

COM: Generally referred to in the literature as M. 
repanda, a different species (see below). 


Mocis antillesia Hampson, 1913 
(Figs. 56, 57) 


ID: Same size and easily confused with M. latipes. 
Ground color pale brownish to pale yellow. Clothing of 
hind legs usually yellowish in this whereas grayish in 
M. latipes. Smaller than M. repanda (see below). 
DIST: Lesser Antilles, Bahamas. 

GUANA: 4 specimens, 1989, 1990. 

BIO: Unknown. 


Mocis repanda (Fabricius, 1794) 
(Figs. 59, 60) 


ID: Larger than the former two species in the genus, 
4.5-5.5 cm wing span. Smaller specimens of this 
species are larger than the largest specimens of both 
M. latipes and M. antillesia. Distinguished from con- 
geners by shape of postmedial lines in both wings: in 
the forewing it is bent inwards after the angle near 
costa, and fades away before tornus, whereas in the 
former two it is straight and reaches tornus; in the 
hindwing it is strongly angled outwards before tornus 
whereas in the others it is straight. 

DIST: Antilles and Guatemala. 

GUANA: 2 specimens, 1987, 1990. 

BIO: Larvae on Mucuna deeringiana (Bort) Merr. [as 
M. megas] (Fabaceae) (Martorell 1976). 

COM: Generally known in the literature as M. megas 
(Guenée), a junior synonym (Poole 1989). Berio 
(1953) clarified the status of M. repanda and illus- 
trated the male genitalia. 


Ophisma tropicalis Guenée, 1852 
(Fig. 61) 


ID: Same size as M. repanda, but with stouter body. 
Extremely variable in pattern and color. Recognized by 
the small, conspicuous white dot on base of forewing. 
DIST: Southern United States, throughout the An- 
tilles, south to Argentina. 

GUANA: 2 specimens, 1987. 

BIO: Larvae on Cupania americana L. (Sapindaceae) 
(Martorell 1976). 


Ophiderinae 


zeta versicolor (Fabricius, 1794) 
(Figs. 62, 63) 


ID: Medium sized, polymorphic species; ground color 
varies from ferrugineous to dark grayish brown. Rec- 
ognized by the pointed forewing and transverse pale 
dash at middle of forewing costa. 

DIST: Throughout the New world tropics from 
Florida to Argentina. 

GUANA: 23 specimens, 1987, 1989, 1990. 

BIO: Larvae on Canavalia (Fabaceae) (Kimball 1965). 
COM: Commonly referred to in the literature by its 
junior synonym, A. repugnalis (Hiibner). 


Metallata absumens (Walker, 1862) 
(Figs. 64-66) 


ID: Medium sized, highly variable species. Ground 
color varies from reddish brown to gray. Similar to E. 
cacata but readily separated by the dark brown head 
and anterior border of thorax, by the nearly rounded 
border of hindwing, and by the filiform antennae in 
both sexes. Some specimens bear a black reniform 
mark at the end of forewing cell. 

DIST: Southern United States, throughout the An- 
tilles, south to Brazil, including Galapagos. 

GUANA: 12 specimens, 1987, 1989, 1990. 

BIO: Unknown. 


Plusiodonta thomae (Guenée, 1852) 
(Fig. 67) 

ID: Medium sized, dark brown species with some 
shinning golden areas on forewing. Recognized by the 
single dentate expansion on middle of dorsum of 
forewing. 
DIST: Described from St. Thomas, considered en- 
demic to the Antilles. 
GUANA: 4 specimens, 1987, 1989. 
BIO: Unknown. 
COM: It is very likely that the continental species, P. 
clavifera (Walker), is conspecific with P. thomae. There 


30 


is some degree of variation in both the Antillean and 
the continental populations, and specimens from both 
regions intergrade into each other. Plusiodonta clav- 
ifera has been reported from Galapagos (Hayes 1975). 


Syllectra erycata (Cramer, 1780) 
(Fig. 68) 


ID: Medium sized, reddish ferrugineous species. 
Forewing with three transverse lines, angled basad 
near costa, and with one or two small, round pale dots 
on outer side of postmedial line. Males distinguished 
by unique shape of antenna, which is uncommonly 
thick throughout its length except for the tip. 

DIST: Florida, throughout the Antilles, south to 
Brazil. 

GUANA: 3 specimens, 1986, 1987. 

BIO: Unknown. 


Litoprosopus puncticosta Hampson, 1926 
(Fig. 69) 


ID: Large, velvet fuscous species. The narrow wings 
and stout body resemble a small sphingid. Readily dis- 
tinguished from other noctuids of same size on the is- 
land by the orbicular black mark at lower edge of hind- 
wing. 

DIST: Haiti, Virgin Islands. 

GUANA: | specimen, 1989. 

BIO: Unknown, however, a close relative, L. futilis 
(Grote & Robinson), has been found boring into the 
flower stalks of Sabal and Serenoa (Arecaceae) (Fer- 
guson et al. 1991). 


Diphthera festiva (Fabricius, 1775) 
(Fig. 70) 


ID: Medium sized, bright yellow, with an elaborate 
pattern of bluish gray lines and three parallel rows of 
dots parallel to external margin. Hindwing dark gray 
with pale cilia. 

DIST: Widespread throughout the New World tropics, 
from Florida to Argentina. 

GUANA: 2 specimens, 1987, 1989. 

BIO: Larvae on Casuarina equisetifolia L. (Casuari- 
naceae), Corchorus hirsutus L. (Tiliaceae), Schrankia 
portoricensis Urb. (Fabaceae), and Waltheria indica L. 
(Sterculiaceae) (Martorell 1976, Torres 1994). Bright 
colored larvae reared in Brazil on Sida sp. (Malvaceae) 
(VOB), acommon weed in disturbed areas on the island. 
COM: Referred to as Noropsis hieroglyphica (Cramer), 
the junior synonym, in older literature. Its pattern may 
be aposematic. One male was tossed towards a gray 
kingbird, Tyrannus dominicensis (Gmelin), who caught 
it in the air, returned to its perch, tried to swallow the 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


moth, then spit it out and cleaned its beak against the 
branch (VOB pers. obs.). 


Gonodonta bidens Geyer, 1832 
(Fig. 71) 


ID: Showy, medium sized moth; cannot be confused 
with any other species on the island. Forewing velvet 
dark brown; basal and postmedial areas paler, crossed 
with waving dark and reddish brown lines. Hindwing 
dark gray with a bright elongate yellow area at middle. 
Head conspicuously white. 

DIST: Florida, throughout the Antilles, south to Ar- 
gentina. 

GUANA: | specimen, 1990. 

BIO: Larvae on Guarea trichilioides L. (Meliaceae), 
Cupania (Sapindaceae) and Diospyrus (Ebenaceae) 
(Todd 1972a). Adults have been reported to damage 
oranges in northern Mexico by piercing ripening fruits 


(Todd 1959). 


Melipotis acontioides (Guenée, 1852) 
(Fig. 72) 

ID: Medium to large sized, 3.5-5 cm wing span, light 
gray species. Hindwings semitranslucent white with a 
broad gray band along external margin not reaching 
lower angle; often with a small gray dot just before 
lower angle. 
DIST: Florida, Antilles, south to Brazil, including 
Galapagos. 
GUANA: 4 specimens, 1989. 
BIO: Larvae on Delonix regia (Bojer ex Hook.) Raf. 
(royal poinciana) and Parkinsonia aculeata L. 
(Fabaceae) (Martorell 1976, Torres 1994). 
COM: Except for M. acontioides, species of Melipotis 
are difficult to distinguish because they look very sim- 
ilar to each other and there is a high degree of varia- 
tion among specimens within each species. Most 
Melipotis species recorded for Guana also occur in 
southern United States and were reviewed by 
Richards (1939) and illustrated in color by Bordelon 
and Knudsen (1999). 

The species of this genus are often the most abun- 
dant moths at lights in dry areas of the New World 
tropics. One of the reasons is that they feed on various 
leguminous plants such as Acacia, Cassia, Prosopis, 
and other species that are abundant in such habitats. 
During certain collecting trips, especially immediately 
after the beginning of rainy season, they came to light 
in such great numbers that the entire sheet was cov- 
ered, making it impossible to collect any other moths. 
On some occasions the lights had to be disconnected 
and collecting discontinued (VOB pers. obs.). 


VOLUME 56, NUMBER I| 


Melipotis fasciolaris (Hiibner, [1831]) 
(Figs. 73, 74) 


ID: Medium to large sized, 3-4.5 cm wing span, vari- 
able species—the most variable species of the genus 
occurring on the island. In some specimens the pat- 
tern is less contrasting while in others the contrast is 
strong. Most specimens can be distinguished from 
those of other species on the island by the antemedial 
oblique, pale fascia of forewing. In M. fasciolaris the fas- 
cia is straight and uniform in width throughout. In some 
specimens the area basad of the fascia is pale olivaceous. 
DIST: Southern United States, Antilles, south to 
Uruguay. 

GUANA: 7 specimens, 1989. 

BIO: Unknown, however Wolcott (1951) noted “Nu- 
merous caterpillars hiding under loose bark of trees of 
Guaiacum officinale L. (Zygophyllaceae), presumably 
after feeding at night on the foliage”. This observation 
should be verified because this tree generally grows to- 
gether with many leguminous species known as food 
plants of other species of Melipotis. 


Melipotis contorta (Guenée, 1852) 
(Fig. 75) 


ID: Same size as larger specimens of M. acontioides 
and M. fasciolaris, but not as variable. Very similar to 
M. famelica with which it shares the white basal area of 
hindwing, and pale head and dorsal area of thorax. 
Easily separated from M. famelica by the irregular pale 
area at the end of cell. In the latter this is nearly 
rounded, whereas in contorta its lower end extends 
broadly towards the external margin. 

DIST: Florida, Antilles. 

GUANA: 4 specimens, 1989, 1990. 

BIO: Unknown. 


Melipotis famelica (Guenée, 1852) 
(Fig. 76) 


ID: Very similar to M. contorta in size and pattem. Color 
pattern not highly variable but sexually dimorphic. Fe- 
males have pattern less contrasting than males. Some 
males have antemedial fascia tinged reddish brown. 
DIST: Southern United States, Antilles, south to 
Venezuela, including Bermuda (Ferguson et al. 1991). 
GUANA: 9 specimens, 1987, 1989, 1990. 

BIO: Larvae on Leucaena latisiliqua (L.) Gillis & 
Stearn (Fabaceae) (Martorell 1976). 


Melipotis ochrodes (Guenée, 1852) 
(Fig. 77, 78) 


ID: Easily confused with M. indomita, a neotropical 
species also recorded from the Greater Antilles but 


31 


not collected on Guana. Highly variable. Basal area of 
hindwing semitranslucent gray, not whitish as in M. 
contorta or M. famelica, or almost dark gray as in M. 
januaris. 

DIST: Antilles, Mexico, south to Brazil. 

GUANA: | specimen, 1989. 

BIO: Larvae on Prosopis juliflora (Sw.) DC. and 
Schrankia portoricensis Urb. (Fabaceae) (Martorell 
1976). 

COM: The specimen illustrated here, identical to the 
one collected on Guana, matches the series at BMNH 
identified as M. ochrodes (type specimen in MNHN, 
Paris, not examined). This species could represent 
only a smaller form of M. indomita. 


Melipotis januaris (Guenée, 1852) 
(Figs. 79, 80) 


ID: On average slightly smaller than other Melipotis 
species on the island; sexually dimorphic. Males have 
forewing with very contrasting, dark brown pattern; fe- 
males little contrasting, reddish brown. Easily distin- 
guished from all other species on the island by almost 
entirely dark gray hindwing. 

DIST: Southern United States, throughout Antilles, 
south to the Guianas and Colombia. 

GUANA: | specimen, 1987. 

BIO: Larvae on Inga laurina (Sw.) Willd. [as I. fagifo- 
lia] (Fabaceae) (Martorell 1976). 


Ascalapha odorata (Linnaeus, 1758) 
(Witch moth, black witch) 


ID: Distinguished by very large size and broad wings 
showing bluish hue. Sexually dimorphic; males black- 
ish gray, females lighter in color, with more contrasting 
pattern, and with three close, parallel, zig-zag, white 
lines crossing the wings. 

DIST: Originally South American, now Pantropical. 
GUANA: 3 specimens, 1982 (J. Lazell), 1988, 1989. 
BIO: Larvae on various leguminous trees, including 
Acacia, Cassia and Piptadenia (Hayward 1969, Hayes 
1975). Comstock (1936), Schreiter (1936) and Bourquin 
(1947) describe its life history and immature stages. 
COM: Illustrated in several works, including Covell 
(1984), Ferguson et al. (1991), Hayes (1975), and Kim- 
ball (1965). 


Epidromia lienaris (Hiibner, 1823) 
(Fig. 81) 
ID: Large, 5 cm wing span; highly variable, gray fus- 
cous species; forewing with conspicuous reniform 
black mark at middle and a postmedial, almost 
straight, pale line. 
DIST: New World tropics. 


GUANA: | specimen, 1990. 

BIO: The larvae (reported as E. pannosa Guenée) 
were found on Psidiwm longipes (O. Berg) McVaugh 
(Myrtaceae), and were fed in the laboratory on P. 
guajava L., Eugenia axillaris (Sw.) Willd. (Myrtaceae), 
Metopium toxiferum (L.) Krug & Urb., and Rhus co- 
pallina L. (Anacardiaceae) (Dickel 1991). 

COM: This is a widespread and highly polymorphic 
species, described more than 10 times (Becker 2001), 
and is commonly known in the literature as E. ze- 
tophora Guenée (Hayes 1975) and E. pannosa (Solis 
1986, Dickel 1991). 


Manbuta pyraliformis (Walker, 1858) 
(Fig. 86) 


ID: Medium sized, gray species. Forewing speckled 
with small black dots and with an oblique postmedial 
yellowish fascia; basal area of this fascia light gray, dis- 
tal area dark gray. Males with pectinate antennae. 
DIST: Florida and Antilles. 

GUANA: | specimen, 1956. 

BIO: Unknown. 

COM: Poole (1989) listed this species under Epidro- 
mia Guenée, however, its genitalia and pectinate an- 
tennae are similar to those of species currently placed 
in Manbuta Walker (Becker 2001). 


Ephyrodes cacata Guenée, 1852 
(Figs. 82, 83) 


ID: Resembling M. absumens in size and coloration 
(see above). Variable in color, from reddish brown to 
gray, mottled with black scales. Distinguished by the 
strongly angled termen of both wings, especially of the 
hindwing, forming a small tail. Male antennae strongly 
pectinate, female filiform. 

DIST: Southern United States, throughout the An- 
tilles, south to Colombia. 

GUANA: 5 specimens, 1989. 

BIO: Larvae on Sesbania grandiflora (L.) Pers. 
(Fabaceae) (Brunner et al. 1975). 


Concana mundissima Walker, [1858] 
(Fig, 84) 


ID: Medium sized, silky shining gray species; forewing 
with fine, broken, transverse lines and a dark dot near 
middle, closer to dorsum. Hindwing semitranslucent 
white, edged with gray. 

DIST: Florida, throughout the Antilles, south to 
Brazil. 

GUANA: 7 specimens, 1989, 1990. 

BIO: Unknown. 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Massala asema Hampson, 1926 
(Fig. 85) 


ID: Medium sized, 3.5 cm wing span, stout bodied, 
pale brownish species. Wings shaded brown with ill 
defined, irregular brownish lines, nearly parallel to 
each other, from costa to dorsum. 

DIST: Antilles. 

GUANA: 2 specimens, 1989. 

BIO: Unknown. 


Lesmone formularis (Geyer, 1837) 
(Figs. 90, 91) 


ID: Medium sized, gray, sexually dimorphic species; 
males have two wide ill-defined dark gray bands across 
the wings; in spread specimens the bands are continu- 
ous, crossing both forewing and hindwing. Females 
lack these bands, however, the edge of the postmedial 
band in the hindwing is replaced by a straight yellow 
fascia running from apex to tornus. 

DIST: Southern United States, throughout the New 
World tropics, including Galapagos. 

GUANA: 3 specimens, 1987, 1989. 

BIO: Larvae on Cassia and Mimosa (Fabaceae). 


Lesmone hinna (Geyer, 1837) 
(Figs. 87-89) 


ID: Same size and similar to L. formularis, but distin- 
guished by the conspicuous round, pale dot on the 
forewing cell. 

DIST: Southern United States, throughout the An- 
tilles, south to Brazil. 

GUANA: 5 specimens, 1989. 

BIO: Unknown. 


Baniana relapsa (Walker, 1858) 
(Figs. 92, 93) 


ID: Small to medium sized, pale ochreous, sexually di- 
morphic species. Male forewing with conspicuous tri- 
angular black patch near base, close to dorsum; post- 
medial area black, fading gradually towards termen. 
Females lack the triangular patch and have the distal 
area lighter gray, resembling P. immunis, but readily 
distinguished by the dark brown anterior edge of tho- 
rax. 

DIST: Restricted to the Antilles. 

GUANA: 18 specimens, 1986, 1987, 1989, 1990. 

BIO: Unknown. 


Eulepidotis addens (Walker, 1858) 
(Fig. 95) 


ID: Small, grayish brown; forewing with three straight 
lines across, the medial and postmedial double, en- 


VOLUME 56, NUMBER | 


closing a conspicuous ochreous band. Hindwing with 
diffuse orbicular mark followed by a short tail on the 
lower part of external margin. 

DIST: Antilles. 

GUANA: 12 specimens, 1987, 1989, 1990. 

BIO: Larvae on Inga vera Willd. (Fabaceae) (Mar- 
torell, 1976). 


Eulepidotis modestula (Herrich-Schiffer, 1869) 
(Fig. 94) 


ID: Small, white tinged yellow species, with lines 
crossing the forewing and a short tail on the hindwing 
similar to those of former species. 

DIST: Antilles. 

GUANA: 1 specimen, 1989. 

BIO: Larvae on Ceiba pentandra (L.) Gaertn. (Bom- 
bacaceae) (Martorell 1976). 


Toxonprucha diffundens (Walker, 1858) 
(Fig. 96) 


ID: Small sized, 1.8-2.2 cm wing span, gray species. 
Highly variable; most specimens have pattern as in 
Kakopoda progenies (see below). 

DIST: Antilles, south to Brazil. 

GUANA: | specimen, 1990. 

BIO: Unknown, however, other species in the genus 
have been reared on Acacia (Fabaceae) (Crumb 
1956). 


Kakopoda progenies (Guenée, 1852) 
(Fig. 97) 


ID: Medium sized, 2.5-3.0 cm wing span; broad 
winged, dark gray moth. Wings crossed with many fine 
waving lines alternating black and pale. Very similar to, 
and easily confused with, T. diffundens (see above), 
but readily separated by size. Smaller specimens of 
this species are always larger than the largest of the lat- 
ter. Males are also separated from T. diffundens by the 
pale brush at the tip of abdomen. 

DIST: Florida, throughout the Antilles to Brazil. 
GUANA: 4 specimens, 1987, 1989. 

BIO: Unknown. 

COM: We consider Kakopoda cincta Smith, 1900, 
new synonym, described from Florida, to represent 
the same species. 


Parachabora abydas (Herrich-Schiffer, [1869]) 
(Fig. 98) 


ID: Small to medium sized, cupreous brown species; 
hindwing semitranslucent white, bordered with a 
wide, diffuse gray area, and veins contrastingly gray. 

DIST: Mexico, throughout the Antilles, south to Brazil. 


33 


GUANA: 3 specimens, 1988, 1989. 
BIO: Unknown. 


Cecharismena abarusalis (Walker, 1859) 
(Fig, 99) 


ID: Small brown species with forewing tinged copper 
and ferrugineous; apex of forewing pointed. Very sim- 
ilar to the following, but easily separated by the 
oblique straight medial line. 

DIST: Florida, throughout Antilles, south to Brazil. 
GUANA: 9 specimens, 1987, 1989. 

BIO: Unknown, however, C. nectarea Moéschler has 
been reared in Puerto Rico on Caperonia palustris 
(L.) A. St.-Hil. (Euphorbiaceae) (Schaus 1940). 


Cecharismena cara Moschler, 1890 
(Fig. 100) 


ID: Same size and easily confused with C. abarusalis. 
Forewing with violet hue. Readily distinguished from 
the former by the medial oblique line curved in C. 
cara and straight in C. abarusalis. 

DIST: Antilles. 

GUANA: 2 specimens, 1989. 

BIO: Unknown (see C. abarusalis). 


Glympis eubolialis (Walker, [1866]) 
(Fig. 101) 


ID: Small gray species with forewing crossed, in the 
middle, with ill defined, straight dark brown band; 
area distad to this band usually darker than basal area. 
Shape, size, and color similar to Bleptina species (be- 
low). Easily distinguished from Bleptina by the porrect 
labial palpi, which are long and upturned in Bleptina 
(see below). 

DIST: Antilles. 

GUANA: 22 specimens, 1989, 1990. 

BIO: Unknown, however, the larvae of G. concors 
were found feeding on Sesbania grandiflora (L.) Pers. 
(Fabaceae) in Puerto Rico (Martorell 1976). 


Herminiinae 


Drepanopalpia lunifera (Butler, 1878), 
new combination 


(Figs. 102, 103) 


ID: Small to medium sized, 1.5-2 em wing span, di- 
morphic, dark fuscous species. Males have very long 
labial palpi thickly covered with long scales, reflexed 
over the head to the posterior margin of thorax. Fe- 
males have long, porrect palpi. Male forewing with a 
reniform, pale mark at end of cell; in females this mark 
is reduced to small dot. 

DIST: Antilles. 


34 


GUANA: 9 specimens, 1987, 1989. 

BIO: Unknown. 

COM: The series in VOB, including specimens from 
Guana Island, Tortola, Puerto Rico and Cuba, was com- 
pared with material in BMNH. Males match the type of 
Drepanopalpia polycyma Hampson 1898, new syn- 
onym, and females match the type of Hypena lunifera 
Butler 1878. D. polycyma is also curated in BMNH as a 
synonym of Mastigophorus latipennis Herrich-Schiiffer. 
There is no specimen of M. latipennis in Coll. Gundlach 
(IES, Havana) where the type material of the Cuban 
species described by Herrich-Schiiffer is supposed to 
be deposited. It is possible that some material is in 
MNHU, Berlin. It is possible that both M. lunifera and 
M. polycyma are junior synonyms of M. latipennis. 


Lascoria orneodalis (Guenée, 1854) 
(Figs. 104, 105) 


ID: Small to medium sized, 1.5-2 cm wing span, dark 
fuscous species, resembling D. lunifera. Males have 
labial palpi as in the former species, but the forewing 
has a strong indentation at middle of external margin. 
Females have long upcurved palpi as in Bleptina. 
DIST: Florida, Antilles. 

GUANA: 3 specimens, 1987. 

BIO: Larvae on tomato leaves (Solanaceae) (Martorell 
1976). 


Bleptina hydrillalis Guenée, 1854 
(Fig. 108) 


ID: Dark brown, 1.8-2.0 em wing span; forewing 
crossed by three, well defined, pale lines: a straight line 
near base, and two sinuate lines, one after the reniform 
mark on cell, the other before external margin. Reni- 
form mark usually pale, but black in some specimens. 
DIST: Southern United States, Central America, Antilles. 
GUANA: 34 specimens, 1986, 1989. 

BIO: Unknown. 


Bleptina caradrinalis Guenée, 1854 
(Figs. 106, 107) 


ID: About same size as B. hydrillalis, pale, variable. In 
those specimens with a dark band across forewing, the 
band is closer to the middle. 

DIST: Southern United States, throughout the An- 
tilles, South to Brazil. 

GUANA: 6 specimens, 1989. 

BIO: Larvae reported on dead leaves (Kimball 1965). 


Bleptina menalcasalis Walker, [1859] 
(Figs. 109, 110) 


ID: Medium sized, 2.2—2.7 cm wing span, pale species. 
Forewing diffusely crossed with ill defined, irregular 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


lines. Distinguished from other noctuids of same size 
by the long, upcurved labial palpi. 

DIST: Antilles, south to Venezuela. 

GUANA: 27 specimens, 1986, 1987, 1989, 1990. 

BIO: Unknown, but likely dead leaves (see B. caradri- 
nalis). 


Bleptina araealis (Hampson, 1901) 
(Figs. 167, 168) 


ID: Very small, 1-1.3 cm wing span, variable, fuscous 
to dark fuscous species. Forewing often with a dark 
gray fascia on basal fourth; area basad to fascia paler 
than rest of wing. Easily distinguished from other 
small species on the island by the very long upcurved 
labial palpi and by three very small, round, pale dots 
on forewing, the first just outside the basal band and 
the two others close together at end of cell. 

DIST: Antilles and Florida (Dickel 1991). The first au- 
thor recently collected one male and one female in 
Mexico: Tamaulipas, E] Encino. 

GUANA: 12 specimens, 1987, 1990. 

BIO: Unknown (see previous species). 


Hypeninae 


Hypena lividalis (Hiibner, 1790) 
(Fig. 111) 


ID: Small, gray; forewing with straight, white post- 
medial line; area basad of line olive, area distad gray. 
DIST: South Palearctic, Pantropical. 

GUANA: 2 specimens, 1989, 1990. 

BIO: Larvae on Urtica and Parietaria spp. (Lédl 
1994:502). 

COM: Ina revision of the genus, Lédl (1994) listed six 
synonyms under H. lividalis, two of them originally de- 
scribed from material collected in the Antilles. 


Hypena minualis (Guenée, 1854) 
(Fig. 166) 


ID: Small, dark gray species with little contrasting 
markings. Forewing with a slightly paler basal area, 
separated from the external dark area by an ill defined 
oblique, dark gray line from near base of costa to mid- 
dle of dorsum; a faint dash near apex. The palpi re- 
semble those of female D. polycyma, but readily sepa- 
rated from the latter by the absence of the pale dot on 
cell. 

DIST: Antilles, south to Brazil. 

GUANA: 3 specimens, 1989. 

BIO: Larvae on Sida rhombifolia L. (Malvaceae) (Fer- 
guson et al. 1991). 


VOLUME 56, NUMBER | 


ARCTIIDAE 
Arctiinae 


Hypercompe simplex (Walker, 1855) 
(Figs. 112, 114) 


ID: Large, white; forewing with outer half translucent, 
basal half with a series of annulate black edged spots. 
Abdomen orange with subdorsal pairs of dark bluish 
dots. Females much larger than males; spots on the 
abdomen white. 

DIST: Puerto Rico, Virgin Islands and Lesser Antilles. 
GUANA: 6 specimens, 1987, 1989. 

BIO: Martorell (1976) lists over a dozen hostplants, in- 
cluding Cedrela (Meliaceae), Cissus (Vitaceae), Erech- 
totes (Compositae), Erythrina (Fabaceae), Ipomaea 
(Convolvulaceae), tomato, banana, beans, guava, egg- 
plant and Solanum torvum Sw. (Solanaceae). 

COM: Commonly found in the literature, including in 
Martorell (1976), as Ecpantheria icasia (Cramer), a 
similar species from continental South America. 


Calidota strigosa (Walker, 1855) 
(Fig. 115) 


ID: Large, gray moth with pink abdomen. Forewing 
with veins darker than ground color, shortly inter- 
rupted with pale. Cannot be confused with any other 
moth from the island. 

DIST: Antilles and Southern United States. 

GUANA: 28 specimens, 1984, 1987, 1989, 1990. 

BIO: Dyar (1901:270) reared this species on Guet- 
tarda elliptica Sw. (Rubiaceae) and described its lar- 
vae; Martorell (1976) lists the same plant as its host in 
Puerto Rico. 

COM: Franclemont (1983) and Watson & Goodger 
(1986) resurrected, respectively, C. laqueata (Edwards 
1887) (type-locality: USA) and C. cubensis (Grote 
[1866]) (type-locality: Cuba) from the synonymy of 
strigosa. We believe they are only geographical forms 
of the same species. 

In the resting position, the adult looks cryptic but 
when touched it opens the wings exposing the bright 
pink abdomen that seems to be aposematic. A speci- 
men was picked from the collecting sheet by a pearly- 
eyed thrasher, Margarops fuscatus (Vieillot), and re- 
jected. The same specimen was placed back on the 
wall where the bird often perched. The same bird 
picked up the moth again and then dropped it again 
(VOB pers. obs.). 


Eupseudosoma involutum (Sepp, [1855]) 
(Fig, 116) 


ID: Medium sized, white moth, with red abdomen. 
Cannot be confused with any other species in the island. 


35 


DIST: Southern United States, throughout Antilles, 
south to Argentina. 

GUANA: 4 specimens, 1987. 

BIO: Larvae on guava (Psidium guineense Sw.), Euge- 
nia, Eucalyptus, and other Myrtaceae. 


Utetheisa ornatrix (Linnaeus, 1758) 
(Fig. 117) 


ID: Medium sized, variable white moth. Most speci- 
mens have the forewing white, tinged pink, and a pink 
costa interrupted regularly by dark gray dots. In other 
specimens most of the white is replaced by pink and 
the wing is crossed with transverse rows of dark gray 
dots. The proportion of gray in the hindwing also 
varies; in some specimens it is restricted to the bor- 
ders, while in others it covers most of the area. 

DIST: Southern United States, throughout the An- 
tilles, south to Argentina. 

GUANA: 5 specimens, 1984, 1988, 1989, 1990. 

BIO: Larvae on various species of Crotalaria 
(Fabaceae). 

COM: Commonly found flying during the day in open, 
disturbed areas where its host plants often grow. Pease 
(1973) discussed the variation of this moth in the Vir- 
gin Islands. 


Utetheisa pulchella (Linnaeus, 1758) 
(Fig. 118) 


ID: Easily confused with the dotted form of U. orna- 
trix (see above); easily separated by the dots on dor- 
sum of thorax. Utetheisa ornatrix has three pairs; U. 
pulchella has only three dots. 

DIST: Africa and Asia, now established in the New 
World tropics, but very rare in collections. The first au- 
thor collected two specimens in Brazil, one at Pipa 
Beach, south of Natal, Rio Grande do Norte, and one 
in Planaltina, DF, near Brasilia, which is the southern 
most record for the species in the New World. 
GUANA: | specimen, 1990. 

BIO: Larvae on Myosotis (Boraginaceae) and grasses 
(Hampson 1901). 


Pericopinae 


Composia credula (Fabricius, 1775) 
(Fig. 113) 


ID: Large, black, with body and wings dotted white; 
forewing with deep red markings on basal half below 
costa. 

DIST: Endemic to the Antilles, this is the only species 
of the genus found on the Puerto Rican Bank. 
GUANA: 10 specimens, 1984, 1986, 1987, 1989. 
BIO: Unknown. Its closest relative, C. fidelissima 


36 


Herrich-Schiaffer, from Cuba and Florida, has been 
reared on Canavalia (Fabaceae), oleander (Nerium), 
and Echites (Apocynaceae) (Kimball 1965). 

COM: Commonly referred to in the literature by its 
junior synonym C. sybaris (Cramer). It is a crepuscu- 
lar moth, commonly found flying along the trails be- 
fore dark. Todd (1982) states it “occurs throughout the 
Greater Antilles, and south to Brazil” but that the 
“continental distribution needs to be studied.” Previ- 
ous authors (Forbes 1930, Bates 1933) considered it to 
be endemic to the Antilles. In the series in USNM 
there is only one non-Antillean specimen, an old spec- 
imen labeled only “Brazil,” which we believe to be 
mislabeled. We are not aware of any other continental 
records for this large and colorful species, which 
would not be overlooked by collectors. 


Ctenuchinae 


Empyreuma pugione (Linnaeus, 1767) 
(Fig. 119) 


ID: Large, wasp-like moth with conspicuous red wings 
and black body. Wings bordered dark gray, and body 
tinged iridescent green. 

DIST: Apparently restricted to the Puerto Rican Bank. 
GUANA: 9 specimens, 1984, 1987, 1989, 1990. 

BIO: Larvae on oleander, Nerium oleander L. (Apocy- 
naceae) (Gundlach 1881). 

COM: Day flying; very likely a Miillerian mimic of 
Pepsis rubra (Drury) (Hymenoptera: Pompilidae) (R. 
Snelling det.), as its larvae feed on a toxic plant. Simi- 
lar to other arctiids, the larvae presumably sequester 
alkaloids from the host plant. Referred to in the older 
literature as E. lichas (Cramer). 


Horama pretus (Cramer, 1777) 
(Fig. 121) 


ID: Large, wasp-like, ochreous brown species, distin- 
guished by the broad white band across base of ab- 
domen. 

DIST: Antilles. 

GUANA: 36 specimens, 1984, 1985, 1987, 1989, 1990. 
BIO: Larvae on Cassine xylocarpa Vent. [as Elaeoden- 
dron xylocarpum|] (Celastraceae) (Wolcott 1951). 
COM: Day flying, commonly seen visiting flowers. Pre- 
sumably a Miillerian mimic of Polistes major Palisot de 
Beavois (Hymenoptera: Vespidae) (R. Snelling det.). 


Horama panthalon (Fabricius, 1793) 
(Fig. 120) 


ID: Similar but smaller than H. pretus. Distinguished 
by pattern on abdomen: only a small white dash across 
base, followed by alternating ochreous and black bands. 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


DIST: Southern United States, throughout the An- 
tilles, south to southern Brazil. 

GUANA: 13 specimens, 1986, 1989, 1990. 

BIO: Unknown. 

COM: Dietz and Duckworth (1976) divided the 
species into three subspecies, assigning the Antillean 
population to the nominal form. Very likely a Miiller- 
ian mimic of Polistes crinitus (Felton) (Hymenoptera: 
Vespidae) (R. Snelling det.). 


Cosmosoma achemon (Fabricius, 1781) 
(Fig. 122) 


ID: Small, wasp-like moth with unique combination of 
colors. Mostly orange with vertex of head and subdor- 
sal sides of abdomen metallic bluish green; forewing 
with translucent areas. 

DIST: Antilles south to Brazil. 

GUANA: 10 specimens, 1989, 1990. 

BIO: Unknown. 


Eunomia colombina (Fabricius, 1793) 
(Fig. 123) 


ID: Medium sized, wasp-like moth with black body 
and translucent wings. Forewing bordered black with 
a dark, red mark at end of cell connected with costa. 
Thorax striped white; abdomen with a carmine red 
band across the base, expanding laterally, followed by 
a white band, partially interrupted dorsally; the rest 
banded with alternating, narrow, red and white lines. 
DIST: Antilles. Hampson (1898) gives also Honduras 
and Brazil, but the material studied by him should be 
checked. 

GUANA: 1 specimen, 1989. 

BIO: “Oruga en las convolvulaceas” (Gundlach 1881). 
COM: Listed as E. columbina, a misspelling, by Wol- 
cott (1951). 


Nyridela chalciope (Hiibner, [1831]) 
(Fig. 124) 


ID: Medium sized, wasp-like, black moth with trans- 
parent wings. Head, thorax dorsally, base of wings, 
legs, and abdomen dorsally and laterally with irides- 
cent blue. Antennae yellow. Forewing bordered black 
with an oblique transverse band from middle of costa 
to tornus. 

DIST: Described from Havana, occurs throughout the 
West Indies and Central America, south to Panama. 
GUANA: | specimen, 1990. 

BIO: Larvae on Cupania americana L. (Sapindaceae) 
(Moschler 1890). 

COM: Some authors regard the Central American 


VOLUME 56, NUMBER | 


population as belonging to a separate species, N. 
xanthocera (Walker). 


Lithosiinae 
Afrida charientisma Dyar, 1913 
(Fig. 169) 


ID: Very small, 8-12 mm wing span, slightly variable 
species. Forewing mostly grayish, slightly tinged with 
green, crossed by alternating, ill defined whitish and 
dark gray bands. Easily recognized by the antemedial 
whitish band across forewing, starting from costa and 
running obliquely outwards to middle, then bent to 
base towards dorsum. 

DIST: Antilles. 

GUANA: 27 specimens, 1985, 1986, 1990. 

BIO: Unknown, however, most of the species of this 
subfamily are lichen-feeders (Hampson 1900). 


Progona pallida (Méschler, 1890) 
(Fig. 170) 


ID: Small, pale species, with no markings. Forewing 
slightly dusted gray. 

DIST: Previously known only from Puerto Rico. 
GUANA: 6 specimens, 1986, 1990. 

BIO: Unknown. 


Lomuna nigripuncta (Hampson, 1900) 
(Fig. 171) 


ID: Small, 1.3-1.7 cm wing span, whitish moth. 
Forewing dusted gray, conspicuously spotted with 
small, dark gray dots. 

DIST: Previously known only from Puerto Rico (Field 
1952). 

GUANA: 29 specimens, 1984, 1985, 1986, 1990, 1991. 
BIO: Unknown. 


GEOMETRIDAE 


Oenochrominae 


Almodes terraria Guenée, [1858] 
(Fig. 134) 


ID: Medium sized, gray species. Wings densely dusted 
with dark gray scales and crossed with pale and dark ill 
defined, irregular bands. Males easily recognized by 
the strongly pectinate antennae nearly the length of 
forewing. Females have a slight olivaceous tinge, and 
external margins of both wings more strongly dentate. 
DIST: Southern United States, throughout the Antilles 
to Colombia. 

GUANA: 2 specimens, 1990. 

BIO: Unknown. 


OW 


Ennominae 


Pero rectisectaria (Herrich-Schiffer, [1855]) 
(Figs. 125, 126) 


ID: Medium sized, sexually dimorphic, variable 
species. Males with ground color varying from pale to 
dark brown; forewing with antemedial band, when 
visible, strongly bent to the base near costa, post- 
medial band nearly straight, with area distad to it 
much paler than rest of wing. Females more reddish 
brown. Easily distinguished from other species of 
same size by the conspicuously scalloped forewing 
margin near apex. 

DIST: Puerto Rico, throughout the Lesser Antilles. 
GUANA: 42 specimens, 1986, 1987, 1988, 1989, 1990. 
BIO: Unknown. 

COM: Poole (1987), who revised this large genus, 
stated that “Either it is rare, or it occurs in areas not 
commonly collected.” Judging from the long series col- 
lected by us, it is not rare. It was common in October 
1989, just after hurricane Hugo, when the first author 
had dozens of specimens on the light and selected 10 
males and 4 females. 


Oxydia vesulia (Cramer, [1779]) 
(Fig, 127) 


ID: Large, extremely variable species, with no two 
identical specimens. Ground color of both wings vary- 
ing from pale yellow, through pale gray to brown, 
clouded and irrorated in various degrees by gray. 
Forewing with an oblique post medial band from apex 
to near middle of dorsum. Easily recognized by the 
white vertex of head. 

DIST: Southern United States, Antilles, south to Ar- 
gentina. 

GUANA: 4 specimens, 1987. 

BIO: Larvae on Acalypha (Euphorbiaceae), Cinchona 
(Rubiaceae), Cissampelos (Menispermaceae), Citrus 
(Rutaceae), Persea (Lauraceae), Rosa (Rosaceae), and Se- 
curidaca (Polygalaceae) (Martorell 1976, Torres 1992). 


Erastria decrepitaria (Hiibner, [1823]) 
(Figs. 128, 129) 


ID: Medium sized, yellow moth. Sexually dimorphic: 
males suffused olive, especially forming a wide bar 
along external margin; females more yellowish with 
the external olive area reduced to a faint irregular 
band, with a conspicuous dark spot on the band near 
the tornus. 

DIST: Southern United States, throughout Antilles, 
south to Brazil. 

GUANA: 4 specimens, 1987, 1989. 

BIO: Unknown. 


38 


Sphacelodes fusilineatus (Walker, 1860), 
revised status 
(Figs. 130, 131) 


ID: Medium sized, dimorphic species. Males dark olive 
brown; forewing crossed with three equidistant, nearly 
parallel lines and with a subtriangular gray mark on costa 
between medial and postmedial line. Females brown 
with lines same as males, but mark on costa absent. 
DIST: Antilles. 

GUANA: 25 specimens, 1986, 1988, 1989, 1990. 

BIO: Unknown. 

COM: This species has been confused in the literature 
with S. vulnerarius (Hiibner). The two differ in many 
features, the most obvious the color of antenna and 
costal mark of forewing. In S. fusilineatus the antenna 
and costal mark are light gray, whereas in vulnerarius 
the antenna is ochreous and the costal mark reddish 
brown. Sphacelodes fusilineatus was described from 
material with no locality label. There is a series in VOB 
collected in Guana, Tortola, St. Thomas, and Cuba. 
One of these was compared with and matches the type 
of S. fusilineatus. It is very likely that this is the species 
referred to by Kimball (1965:192) as “5221, 1S. SP.”, 
and listed by Ferguson (1983:95) as S. haitiaria 
Oberthiir. The series of S. vulnerarius in VOB was col- 
lected in Cuba, Mexico, and Brazil. In Cuba, S. fusilin- 
eatus was captured at Pinares de Mayari, Holguin, 
while those of vulnerarius at Vinales, Pinar del Rio. 
The first locality is dry, similar to the conditions in St. 
Thomas and Guana, while the second is humid, similar 
to those areas on the American continent from where 
the series of S. vulnerarius came. 


Macaria paleolata (Guenée, [1858]) 
(Fig. 132) 

ID: Small, slightly variable, pale species. Wings 
crossed by two ill defined, nearly straight bands; area 
distad to postmedial band darker than rest of the wing. 
Distinguished by head and anterior margin of thorax 
ochreous. 
DIST: Antilles. 
GUANA: 24 specimens, 1989, 1990. 
BIO: Unknown, however, other species of this large 
genus feed on leguminous species (Fabaceae). 
COM: This species was transferred from Semiothisa to 
Macaria by Scoble (1999). It is very likely that other, 
similar species of the large genus Macaria occur on the 
island. 


Patalene ephyrata (Guenée, [1858]) 
(Fig. 133) 
ID: Small, variable, pale yellow to pale brown species. 
Forewing with antemedial band nearly evenly 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


rounded, postmedial band straight to near apex then 
strongly angled basad to costa. Hindwing with a single, 
straight band near middle. Distinguished by pointed 
apex and the sinuate external margin of forewing. 
DIST: Antilles (Herbulot 1984). 

GUANA: 41 specimens, 1986, 1987, 1988, 1989, 1990. 
BIO: Larvae on Ficus (Moraceae) and Ricinus (Eu- 
phorbiaceae) (Brunner et al. 1975). 


Cyclomia mopsaria Guenée, [1858] 
(Figs. 182, 183) 


ID: Small, extremely variable species. Ground color of- 
ten pale tinged reddish brown, to dark reddish brown. 
Forewing varying from unmarked to marked with 
transverse, ill defined, irregular lines. Hindwing pale to 
orange, often bordered with reddish brown. Similar to 
small noctuids such as Eublemma spp., but distinguished 
by the bipectinate antenna in males. Distinguished from 
other small geometrids by sharply pointed, porrect 
labial palpi, unusually long for a geometrid. 

DIST: Antilles, south to Brazil. 

GUANA: 11 specimens, 1987, 1989, 1990. 

BIO: Unknown, however, Erythroxylum havanense 
Jacq. (Erythroxylaceae) has been listed as food plant of 
“Cyllomia sp. [presumably a misspelling] Un gusano 
medidor,” in Cuba (Brunner et al. 1975). 

COM: Its extreme variation led to its description sev- 
eral times. It is likely that after revision more names 
will be added to the six junior synonyms currently 
listed (Becker in prep.). 


Geometrinae 


Eueana simplaria Herbulot, 1986 
(Figs. 12-14, 141, 142) 


ID: Small to medium sized, bluish green species. 
Lines on wings almost indistinct. Vertex of head white; 
abdomen with traces of a whitish line dorsally. Geni- 
talia (Figs. 12-14) very similar to those of E. niveoci- 
liaria (Herrich-Schiffer), illustrated in Ferguson 
(1985: fig. 25a-e). 

DIST: Guadaloupe. 

GUANA: 17 specimens, 1986, 1989. 

BIO: Unknown, but Scoble (1999) records E. niveocil- 
iaria from Rhamnaceae. 

COM: Of all “greens” from the island this is the only 
bluish species, and it is slightly larger than the other 
species. The subfamily was revised for North America by 
Ferguson (1985) and for the neotropics by Pitkin (1996). 


Phrudocentra centrifugarium (Herrich Schiffer, 1870) 
(Figs. 143-146) 


ID: Medium sized, extremely variable, bright green 
species. Some specimens, usually males, have only 


VOLUME 56, NUMBER 1 


small, dark brown dots on wings, one on cell and the 
others along what would be the antemedial and post- 
medial lines; others have larger marks, of various sizes 
and shapes, whitish or brownish, above tornus. Ab- 
domen has a series of minute, white dots dorsally, one 
on each segment. 

DIST: Florida, throughout Greater Antilles to Puerto 
Rico (Ferguson 1985). 

GUANA: 4 specimens, 1990. 

BIO: Larvae on Myrica cerifera L. (Myricaceae) 
(Scoble 1999). 


Chloropteryx paularia (Méschler, 1886) 
(Fig. 186) 


ID: Small, olivaceous species. Wings with antemedial 
and postmedial rows of minute whitish dots forming 
irregular lines. This is the only species of the green 
Geometrinae with this color. 

DIST: Florida, throughout the Antilles. 

GUANA: 11 specimens, 1987, 1989, 1990. 

BIO: Larvae on Myrica cerifera L. (Myricaceae) in 
Florida (Ferguson 1985). 


Synchlora frondaria (Guenée, [1858]) 
(Fig. 187) 


ID: Small, bright green species. Distinguished from S. 
cupedinaria by the concolorous fringes. 

DIST: United States, throughout the Antilles, south to 
Argentina. 

GUANA: 29 specimens, 1986, 1988, 1989, 1990. 

BIO: Larvae on Stilingia (Euphorbiaceae), Pluchea, 
Bidens, Chrysanthemum, Helianthus (Asteraceae), 
Rubus (Rosaceae), Glycine max (L.) Merr., and 
Prosopis (Fabaceae) (Ferguson 1985). 


Synchlora cupedinaria (Grote, 1880) 
(Fig. 188) 


ID: Small, bright green species. Wings thickly bor- 
dered with brown. Thorax and abdomen brown dor- 
sally; abdomen usually with white dots dorsally. Easily 
distinguished from the other green Geometrinae by 
the brown thorax and abdomen. 

DIST: Florida, throughout Greater Antilles to Virgin 
Islands, to Nevis (Herbulot 1984). 

GUANA: 41 specimens, 1985, 1986, 1988, 1989, 1990. 
BIO: Larvae on Lantana camara L. (Verbenaceae) 
(Scoble 1999). 


Sterrhinae 


Of all the moths treated in this paper, the Sterrhinae 
have proven to be most difficult to identify. We have 
tried to reflect the species concepts as represented in the 
BMNH and USNM collections, and have followed the 


39 


generic placements in Scoble (1999). Our dissections of 
types, as well as Guana specimens, indicate that many 
problems exist in the existing classification, and full reso- 
lution of the names is beyond the scope of this paper. 


Semaeopus malefidarius (Méschler, 1890) 
(Fig. 135) 


ID: Medium sized, pale moth densely irrorated with 
reddish brown scales. Wings crossed with ill defined, 
hardly contrasting, irregular bands, slightly darker than 
ground color. 

DIST: Puerto Rico, Virgin Islands. 

GUANA: 14 specimens, 1989, 1990. 

BIO: Unknown. 

COM: Closely related to S. castarium (Guenée) from 
the Greater Antilles. 


Leptostales noctuata (Guenée, | 1858]) 
(Figs. 136, 137) 


ID: Small to medium sized, pale species, densely irro- 
rated with olive scales. Forewing with antemedial and 
postmedial bands darker than ground color, termen 
sinuate, apex pointed. Males show a dark mark on cell; 
in females this mark is reduced and faint. 

DIST: Antilles. 

GUANA: 19 specimens, 1986, 1987, 1988, 1989, 1990. 
BIO: Unknown. 


Acratodes suavata (Hulst, 1900) 
(Fig. 185) 


ID: Small, white species; wings crossed with three ill 
defined, little contrasting, grayish bands; medial and 
postmedial closer together. The only white geometrid 
on the island with no contrasting marks on wings. 
DIST: Southern United States, Antilles. 

GUANA: 31 specimens, 1985, 1986, 1987, 1988, 1989, 
1990. 

BIO: Larvae on Randia aculeata L. (Rubiaceae) (Kim- 
ball 1965). 

COM: It is very likely that A. virgotus (Schaus), de- 
scribed from Jamaica, is a synonym. Our series 
matches the type of A. virgotus in USNM. 


Lobocleta nymphidiata (Guenée, [1858]) 


ID: Small, white species with wings crossed with 4-5 
narrow, irregular dark brown lines, consisting of rows 
of dark dots. 

DIST: Antilles. 

GUANA: | specimen, 1985. 

BIO: Unknown. 


40 


Scopula laresaria Schaus, 1940 
(Fig. 178) 


ID: Small, pale whitish, irrorated with scattered gray 
scales; forewing crossed with poorly defined, irregular, 
narrow pale yellowish lines. Same size and color as L. 
nymphidiata, but readily distinguished by the blackish 
vertex of head, white in L. nymphidiata. 

DIST: Puerto Rican bank. 

GUANA: 9 specimens, 1985, 1988, 1989. 

BIO: Unknown. 


Idaea sp., probably I. fernaria (Schaus, 1940), 
new combination 
(Figs. 179, 180) 


ID: Small, light gray, densely irrorated with dark gray 
scales; wings crossed with three, ill defined, irregular, 
narrow, dark gray lines more or less interrupted, giving 
the impression that they are densely dotted; both 
wings with black dot on cell. Females slightly lighter 
than males. 

DIST: Antilles. 

GUANA: 27 specimens, 1985, 1986, 1988, 1987, 1989. 
BIO: Unknown. 

COM: Based on the structure of male genitalia and 
eigth sternite of a male paratype, this species does not 
belong in Scopula, so we are provisionally transferring 
it to Idaea fernaria, new combination, following the 
generic concept of Holloway (1997). The Guana spec- 
imens are very similar to the type series of Idaea 
fernaria, but differ in the size of the long expansion on 
the juxta. In Guana specimens it is much longer than 
in a paratype of I. fernaria, nearly the size of the val- 
vae, expanding beyond the uncus. Idaea fernaria is ex- 
ternally similar to I. amnesta (Prout, 1922), from Ja- 
maica, but the male genitalia are very different. 


“Idaea” monata (Forbes in Ramos, [1947]) 
(Figs. 172, 173) 


ID: Very small, gray species; antemedial and post- 
medial lines dark gray, strongly contrasting, especially 
in the males; in some males the area between the two 
lines are dusted dark gray, forming a wide band across 
the wings. 

DIST: Antilles. 

GUANA: 29 specimens, 1985, 1986, 1989, 1988, 1990. 
BIO: Unknown. 

COM: This species is very similar to Idaea insulensis 
(Rindge, 1958), from Florida, but differs in the base of 
the valvae being more expanded in I. insulensis than in 
I. monata, although study of extensive series could 
show this to be interspecific variation. Based on the 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


structure of the male genitalia (Rindge 1958:fig. 9), 
neither I. monata or I. insulensis are properly placed 
in Idaea (cf. Holloway 1997), but recognition of the 
proper generic placement is beyond the scope of this 
paper. Weakly patterned individuals of I. monata are 
similar externally to Lobocleata nataria (Walker, 1866), 
but the male genitalia do not match the type of L. 
nataria in BMNH. 


Idaea minuta (Schaus, 1901) 
(Figs. 176, 177) 

ID: Very small, 8-10 mm wing span, pale species. 
Wings crossed with several, narrow, ill defined, irregu- 
lar, light brown lines. One of the smallest geometrid 
species on the island. 
DIST: Antilles, USA. 
GUANA: 19 specimens, 1985, 1987, 1988, 1989. 
BIO: Unknown. 
COM: Specimens belonging to this series were identi- 
fied as I. minuta by C. Covell, and also matched the 
type of Ptychopoda curtaria Warren 1904, new syn- 
onym, from Jamaica, in BMNH. 


Idaea eupitheciata (Guenée, [1858]) 
(Figs. 174, 175) 


ID: Very small, sexually dimorphic, pale species, with 
wings crossed with irregular reddish brown bands. 
Males paler than females; forewing with small tuft of 
dark scales near dorsum; hindwing with a small tail 
near tornus. Females darker than males and without 
the tufts and tails. 

DIST: Antilles. 

GUANA: 16 specimens, 1985, 1988, 1989, 1990. 

BIO: Unknown. 


Leptostales phorcaria (Guenée, [1858]) 
(Fig. 181) 


ID: Small, reddish brown; wings crossed with narrow, 
ill defined, irregular, yellowish lines; thorax dorsally 
and forewing costa yellow. Cannot be confused with 
any other species on the island. 

DIST: Antilles. 

GUANA: 2 specimens, 1989. 

BIO: Unknown. 


Leptostales oblinataria Méschler, 1890 
(Fig. 184) 


ID: Small, olivaceous species, with a broad reddish 
brown band across the forewing, delimited by medial 
and postmedial bands. Some specimens with this band 
faded. Cannot be confused with any other species on 


the island. 


VOLUME 56, NUMBER 1 


DIST: Southern United States, Antilles, into South 
America (Covell 1969). 

GUANA: 15 specimens, 1988, 1989, 1990. 

BIO: Unknown. 


Larentiinae 


Obila praecurraria (Méschler, 1890) 
(Fig. 138) 


ID: Large species with velvet moss green forewing and 
orange ochreous hindwing. Forewing crossed with nu- 
merous alternating pale and dark gray waving bands. 
Hindwing with a wide gray band extending inwards, 
along internal margin, to the base. Female with middle 
of forewing crossed with a wide, irregular whitish band. 
DIST: Antilles. 
GUANA: 1| specimen, 1989. 

_ BIO: Unknown. 
COM: Our specimen matches the type of Pterocypha 
xantholiva Warren, 1895, synonymized with this by 
Schaus (1940:326). 


Pterocypha defensata Walker, 1862, revised status 
(Figs. 139, 140) 


ID: Medium sized, variable, moss gray. Similar to O. 
praecurraria, but slightly smaller and lacking the or- 
ange ochreous hindwing color. 

DIST: Southem United States, Antilles. 

GUANA: 20 specimens, 1986, 1987, 1989, 1990. 

BIO: Unknown. 

COM: Commonly referred to in the literature, includ- 
ing Scoble (1999), by its synonym P. floridata (Walker), 
but P. defensata was described in 1862 and P. floridata 
in 1863, so defensata has priority. 


HYBLAEIDAE 


Hyblaea puera (Cramer, 1777) 
(Fig. 150) 


ID: Medium sized, fuscous species with hindwing 
beautifully decorated with yellow to orange patches. 
Abdomen dorsally crossed with narrow lines behind 
each segment. 
DIST: Pantropical. 
GUANA: 2 specimens, 1989, 1990. 
BIO: Larvae on Crescentia cujete L., Spatodea cam- 
panulata P. Beauv., Tabebuia heterophylla (DC.) Brit- 
ton (Bignoniaceae), and Petitia domingensis Jacq. 
(Verbenaceae) (Martorell 1976). Considered as a mi- 
nor pest of forest trees in Puerto Rico (Torres 1994). 
Immature stages described by Singh (1995). 

COM: Taxonomy of the related species discussed by 
Berio (1967), but H. puera apparently represents a 
species complex (Shaffer & Nielsen 1996). 


41 


COSSIDAE 


Psychonoctua personalis Grote, 1865 
(Figs. 147-149) 


ID: Medium to large, 2-4.5 cm wing span, narrow 
winged, gray species. The short, strongly pectinate an- 
tennae, and abdomen thickly clothed with long scales 
makes it easily recognized from any other large moths 
on the island. 

DIST: Antilles and Mexico. 

GUANA: 5 specimens, 1990. 

BIO: The larvae are wood borers in many trees, some- 
times causing severe damage to orange, coffee, white 
mangle, sea grape, etc. (Wolcott 1951). 


ACKNOWLEDGMENTS 


Sampling on Guana Island was supported by The Conservation 
Agency, through a grant from the Faleconwood Corporation. We 
thank James Lazell for providing arrangements to work on the is- 
land. George Proctor, formerly of the Puerto Rico Department of 
Natural Resources, provided information on plants. Research facili- 
ties were provided by the National Museum of Natural History, 
Smithsonian Institution and the Natural History Museum, London. 
We thank Robert W. Poole and Douglas C. Ferguson (both formerly 
U.S. Dept. Agriculture Systematic Entomology Laboratory), Martin 
R. Honey (BMNH), Charles Covell (University of Louisville, Ken- 
tucky), Jean-Marie Cadiou, John Wilterding and Morton S. Adams 
for assistance with moth identifications. Poole also allowed us to in- 
clude his generic description in the appendix. Roy Snelling (Los An- 
geles County Museum) provided identifications of wasps. Liao Wei- 
Ping identified the birds. The plates were photographed by Chip 
Clark (Smithsonian), with assistance from Ronald W. Hodges 
(USDA). John Brown and Michael Pogue reviewed the manuscript. 
Karolyn Darrow and Maia Vaswani provided assistance at USNM 
and BMNH, respectively. The line drawings of genitalia were pre- 
pared by W. Cavalcanti, EMBRAPA-CPAC. Most of the work on 
this paper was done while Becker worked for EMBRAPA-CPAC, 
Planaltina, and Miller for Bishop Museum, Honolulu. 


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GuNDLACcH, J. 1881. Contribucion a la Entomologia cubana. G. 
Monteil, Havana. 445 pp. 

HAILE, D. G., J. W. Snow & J. R. YouNG. 1975. Movement by adult 
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4:295-226. 

Hampson, G. F. 1898. Catalogue of the Lepidoptera Phalaenidae 
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. 1900. Catalogue of the Lepidoptera Phalaenidae in the 

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. 1905. Catalogue of the Lepidoptera Phalaenidae in the 

British Museum. Vol. 5. 634 pp. 

1910. Catalogue of the Lepidoptera Phalaenidae in the 
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Hayes, A. H. 1975. The larger moths of the Galapagos Islands 
(Geometroidea, Sphingoidea & Noctuoidea). Proc. Calif. Acad. 
Sci. (4) 40:145-208. 

Haywarb, K. J. 1969. Datos para el estudio de la ontogenia de lep- 
id6pteros argentinos. Tucuman, Fundacion e Instituto Miguel 
Lillo. Miscelanea no. 31. 142 pp. 

HEATWOLE, H., R. LEviNs & M. D. Byer. 1981. Biogeography of 
the Puerto Rican Bank. Atoll Res. Bull. 251:1—55. 

HERBULOT, C. 1984. Geometridae de Nevis (Lepidoptera). Bul- 
letin de la Societe Entomologique de Mulhouse 1984:27-29. 

Hopces, R. W. 1971. Sphingoidea. In Dominick, R. B. et al. (eds.), 
The moths of America north of Mexico. Fasc. 21. E. W. Classey 
Ltd. and R. B. D. Pub., Inc., London. 158 pp. 

Ho.ioway, J. D. 1996. The Lepidoptera of Norfolk Island, actual and 
potential, their origins and dynamics, pp. 123-151. In A. Keast & 
S. E. Miller (eds.), The origin and evolution of Pacific Island bio- 
tas, New Guinea to eemiann Polynesia: patterns and processes. 
SPB Academic Publishing by, Amsterdam. vi + 531 pp. 

. 1997. The moths of Borneo: Family Geometridae, Sub- 
families Sterrhinae and Larentiinae. Malayan Nat. J. 51:1-242, 

Il, =n, 

Honor AY, J. D., G. Kippy, D. PEGGIE, D. J. CarTER & S. E. 
MILLER. 2001. Families of Malesian moths and butterflies. 
Brill, Leiden. xi + 455 pp. 

Ho.ioway, J. D. & E. S. NIELSEN. 1998. Biogeography of the Lep- 
idoptera, pp. 423-462. In N. P. Kristensen (ed.), Lepidoptera, 
moths and butterflies. Vol. 1: evolution, systematics, and bio- 
geography. Walter de Gruyter, Berlin and New York. 

KiMBALL, C. P. 1965. Lepidoptera of Florida. Gainesville, Florida 
Dept. Agriculture. 363 pp. 

KiRIAKOFfF, S. G. 1976. Agaristidae III (American genera). Das 
Tierreich 99:i-vi, 1-86 (“1977”). 

Kitcuine, I. J. & J.-M. Capiou. 2000. Hawkmoths of the world: an 
annotated and illustrated revisionary checklist (Lepidoptera: 
Sphingidae). Cornell University Press, Ithaca. viii + 227 pp: 

KRISTENSEN, N. P. (ED.). 1998. Lepidoptera, moths and butterflies. 
Vol. 1: Evolution, systematics, and biogeography. Walter de 
Gruyter, Berlin and New York. x + 491 pp. 


VOLUME 56, NUMBER 1 


LAFONTAINE, J. D & R. W. PooLe. 1991. Noctuoidea, Noctuidae 
(Part), Plusiinae. In Dominick, R. B. et al. (eds.), The moths of 
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LODL, M. 1994. Revision der Gattung Hypena Schranck, 1802 s.1., 
der iithiopischen und madagassischen Region, Teil 1. Ann. 
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MILLER, J. Y. 1994. Behavior in butterflies as a means of conserva- 
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PEASE, R. W., JR. 1973. Variation of Utetheisa ornatrix (Arctiidae) 
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POOLE, R. W. 1989. Noctuidae. Lepid. Cat. (new series) 118:1-1313. 

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43 


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of Anegada, British Virgin Islands, with ere of a new 
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1972b. A note on the generic transfer of “Catabena” esula 

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Received for publication 12 July 2001; revised and accepted 18 
November 2001. 


44 


APPENDIX 
Catabenoides Poole, new genus 


[The following new generic description was in- 
tended to appear in a fascicle of the series Moths of 
America North of Mexico. Because this fascicle will not 
be published in the foreseeable future, the author has 
permitted us include the new generic name here, in 
order to allow us to describe Catabenoides lazelli in 
the proper place. ] 

Type-species: Adipsophanes terminellus Grote, 
1883 

Catabenoides contains the majority of the species 
previously placed in Catabena. The genus appears to 
be an outlier of a large group in southern South 
America, primarily Paraguay and Argentina. The 
genus is postulated to be closest phylogentically to 
Catabena and Neogalea, but it has a number of curious 
synapomorphies making its affinities somewhat prob- 
lematical. The principle identifying characters are in 
the male and female genitalia. In the male valve the 
bottom margin of the sacculus in both the right and 
left valves is produced into a long process separate 
from the clasper. The ventral margins of the valvae 
have strong elongate setae. The sacculus is lightly chi- 
tinized proximal to the clasper separating off a much 
more heavily sclerotized plate. 

In addition to the three species known from North 
America (the type species, C. vitrina (Walker), new 
combination, and C. divisa (Herrich-Schiffer), new 
combination), there is one previously described 
species; Catabenoides seorsa (Todd) (Catabena seorsa 
Todd), new combination, from the Galapagos Is- 
lands. There are two undescribed species in the West 
Indies [one of these is described herein as C. lazelli], 
one undescribed species from central Mexico, and at 
least two unnamed species from Paraguay. The exact 
affinities of Catabenoides, Catabena, and Neogalea 
with the Argentina and Chilean faunas remain to be 
determined. The single remaining described species in 
“Catabena” of Poole (1989), Laphygma terens Walker, 
was described from “Venezuela.” The type is rubbed 
and its abdomen is missing. I have not been able to 
match it with any specimens from the extensive 
Venezuela material in the USNM. It appears superfi- 
cially to be a Catabenoides. Therefore for book-keeping 
purposes, I place it as Catabenoides terens (Walker), 
new combination. 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Description. Head: Lashes absent; eyes large, 
naked; frons slightly swollen, very closely scaled; an- 
tenna simple, faintly ciliate; palpi unremarkable for the 
tribe; proboscis normal; two thin ridges of flattened 
scales between the bases of the antennae. Thorax: 
Patagia capable of being raised in a hood; vestiture of 
dorsum of thorax of flattened scales without hairs; 
slight metathoracic tuft; vestiture of wings of flattened 
scales; no sign of sexually modified scales in male. Pro- 
thoracic leg: Tibia with later ridge of hair giving it a 
flattened look; no tibial claw; tibia approximately as 
long as first three tarsal segments; first four tarsal seg- 
ments with three rows of spines; tarsal claw without a 
tooth. Mesothoracic and metathoracic legs: Proxi- 
mal third of tibia with tuft of long hairs and scales; tibia 
approximately as long as first three tarsal segments. 
External tympanic region: Approximately as in Supra- 
lathosea Barnes & Benjamin, however, first tergum much 
shorter than in Catabena and Supralathosea and with a 
strong definite proximal lip as in Apharetra Grote; hood 
strong. Internal tympanic region: Not examined. 
Abdomen: Very weak tuft on first tergite; male with 
basal hair pencils and accessory hair pencils in known 
species. Eighth sternum with strong row of hairs in 
middle of U-shaped pleurite. Male genitalia (Figs. 2, 
3): Valvae characterized by production of ventral mar- 
gin of sacculus into a process of various shapes; ventral 
margin of valvae with strong modified setae; a weak un- 
sclerotized area before origin of clasper; uncus swollen 
with an apical tooth; juxta a single pointed plate; vesica 
in type species elongate-ovate, connected by a short 
neck to body of aedeagus; type species with a group of 
elongate, fused spines at apex of vesica and with 
groups of short, stubby spines near the middle and 
ventral margin of the vesica; vesica variable in other 
species. Female genitalia: Ovipositor lobes square, 
unmodified; ductus bursa well sclerotized, elongate, 
mushroom shaped in type species, separated by a dis- 
tinct junction from the heavily sclerotized upper part of 
bursa; bursa bilobed, but lobes not distinctly separate; 
corpus bursae with stellate ridges, but no signum. Larva 
and foodplants: Unknown for North American species 
but recorded as Lantana peduncularis Andersson (Ver- 
benaceae) in Catabenoides seorsa from the Galapagos. 


Robert W. Poole 
Nearctica 
Rockville, Maryland 


BOOK REVIEWS 


Journal of the Lepidopterists’ Societ 
56(1), 2002, 45-46 


THE FAMILIES OF MALESIAN MOTHS AND BUTTER- 
FLIES, by J. D. Holloway, G. Kibby and D. Peggie, with 
contributions by D. Carter and S. E. Miller and photo- 
graphic color plates by B. D’Abrera. 2001. Brill Aca- 
demic Publishers, Leiden, Netherlands. ISBN 90-04- 
11846-2. List Price: U.S. $124, EUR 118, NLG 222.57 


This important treatment of the Lepidoptera is part 
three of the Fauna Melesiana Handbook series. For 
those wanting the basic systematics, morphology and 
biology, and to identify families of butterflies and 
moths quickly and easily, this is the handbook for you. 
But it is much more than that. The scholarship and 
writing style provide historical and current perspec- 
tives of where interesting research questions lie. Al- 
though expensive, this well produced, solidly bound 
book will remain an important resource for research 
on Lepidoptera for many years to come. 

Chapter 1 introduces principles and practices of 
classification, phylogenetic relationships, diversity, and 
biogeography of Malesian Lepidoptera. The tone is a 
balance of engaging enthusiasm for the Lepidoptera 
and a rigorous understanding of their systematics that 
is at a gentle simmer rather than a furious boil. Al- 
though the systematic techniques advocated utilize 
strict methods, they are elegantly interjected such that 
the reader is drawn in and convinced of their impor- 
tance rather than bludgeoned in fundamentalist man- 
ner as in many modern treatments. This chapter also 
includes a useful section (contributed by D. Carter) on 
the practicalities of making, studying and managing 
collections. 

Chapter 2 is a uniformly excellent summary of early 
stage and adult morphology supported with lucid, an- 
notated illustrations. It demystifies the morphology of 
Lepidoptera such that one is eager to have a look for 
oneself. As such Chapter 2 is certain to be consulted 
repeatedly by veterans wanting comparative facts and 
reminders, and because of its clarity it will also inspire 
up-and-coming generations of lepidopterists. The di- 
rect, simple style of enthusiasm is once again more en- 
gaging than the obfuscated technicalities given in 
many texts of this type. 

Chapter 3 is concerned with identification. Rather 
than simply presenting a key that is logical only to 
those who wrote it, the authors give a short review of 
previous identification keys pertinent to the fauna, and 
provide cautions for the beginning lepidopterist (or re- 
minders to the veterans) that few specimens that come 
in for examination are “perfect” —perfection being an 


implicit assumption in all keys. The authors engender 
a suspicion that one must develop an appreciation for 
the variation in specimen condition, and then through 
practice, a familiarity with the major groups, and even- 
tually experience will aid the user in identifying the 
specimen at hand. In essence the authors acknowledge 
that mistakes are normal in the course of learning, and 
that one must learn to read between the lines of spec- 
imens, so to speak. To this end we are shown that cer- 
tain characters in the key are important: venation, tym- 
bal organs, head structures, and even resting postures 
of the live insects, but they may show variation, and 
this is interesting. Proceed to the keys with confidence. 
Here we find a no-nonsense approach that not only in- 
cludes keys to adults, but subsequent treatments of 
larvae also point to important characters (not just tech- 
nical ones) that easily separate various groups: prolegs, 
spines, hairs, filaments and beyond. Finally we are 
provided a summary of the foods caterpillars in major 
groups feed upon (leaves, bark, wood, roots, fungus, 
other insects, etc.), and the major themes of host plant 
associations in the various families. 

Chapter 4 provides the banquet: detailed accounts 
of each family that offer pertinent taxonomic, behav- 
ioral and ecological characteristics for each group. The 
experience distilled over a lifetime study of the Lepi- 
doptera supplies the rhythms, melody, tenor and taste- 
ful improvisation. The accounts are extremely well 
done, and here I seem to detect Holloway’s literary 
stamp—an inherent mixture of classical scholarship 
and awareness of new horizons. 

We are offered eight color plates that are uniformly 
excellent. However, given the thoroughness and scope 
of the text (and the monetary price), one immediately 
asks the question, why so few? Whatever the con- 
straints were during production it is unfortunate that 
this book was not allocated many more color plates. An 
increased number of plates would have gained a wider 
audience and amplified the accessibility of an already 
impressive work. 

There are two appendices. The first compares 
species richness of families among relevant areas, and 
puts them into context of the world fauna. Those in- 
terested in comparative diversity will use this section. 
The second appendix is a comprehensively detailed 
tabulation of known pest species, thus providing an 
important resource for those with a leaning toward the 
agricultural. 

And then there is the reference section. Admirable. 
Wonderfully rich. Redolent of quiet, vertical tasting 
within the cellars of classical scholars. One could, and 
should get lost within this section for years to come. 
What more can I say? 


46 


The two indexes, one to morphological and the sec- 
ond to the scientific names, are both equally good. 
However, confining the indexes only to these topics 
does omit some things in the behavioral and evolution- 
ary realm that would be of interest to many users. For 
example, a favorite topic of mine, myrmecophily, is not 
in either index. Rather, the trait must be sought via 
browsing with prior suspicion in particular groups 
(some Lycaenidae and Riodinidae). But the fact that 
myrmecophily also occurs in other, less well-known 
groups (e.g., Cyclotornidae) cannot be accessed via the 
index, it must be stumbled upon. A general index 
would have been welcome and useful indeed. 

It may be one of those humorous examples where 
British and Americans are ‘divided by a common lan- 
guage’, but I was puzzled by one word choice. In this 
book the term used to denote the need for experience 
to develop a “gestalt” to identify various groups has re- 
peatedly and amusingly sprung unbidden to my mind. 
The fact that experience imparts the ability to cor- 
rectly identify butterflies and moths by a “feel” is a 
very real phenomenon among experts. But using the 
word “jizz” to describe the result of that mental process 
(and including it in the index) conjures up dated, but 
seminal anene an hipster slang that typically means 
something altogether different. What an interesting ex- 
periment in linguistic evolution has been set in motion! 

In sum, I believe this important handbook will in- 
spire genuine enthusiasm for a hands-on research ap- 
proach to topics concerned with Lepidoptera. We 
need more books of this type to counter the facile 

“look but don’t touch” method of studying the natural 
world that is advocated by the popular media, and the 
authors are to be congr: atulated on their ability to rise 
above this. The serious students of Lepidoptera (cur- 
rent and future) who will make fundamental contribu- 
tions toward understanding the natural world will 
thank the authors for this handbook. 


P. J. DEVrIES, Center for Biodiversity Studies, Mil- 
waukee Public Museum, 800 West Wells St., Milwau- 
kee, Wisconsin 53233, USA. 


Journal of the Lepidopterists’ Society 
56(1), 2002, 46-47 


THE GEOMETRID MOTHS OF EUROPE, VOLUME 1, IN- 
TRODUCTION, ARCHIEARINAE, ORTHOSTIXINAE, DESMO- 
BATHRINAE, ALSOPHILINAE, GEOMETRINAE, by Axel 
Hausmann. 2001. Apollo Books. Stenstrup, Denmark. 
282 pp., 8 color plates. 17 cm by 24 cm. ISBN: 87- 
88757-35-8. Retail cost DKK 490,00 (approximately 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


USD 61.00). Hard cover. A 10% discount is offered to 
subscribers of the series when ordered from Apollo 
Books. 


Before the 1968 reprint of Holland’s Moth Book by 
Dover Press, it was not easy for many average persons 
in the U.S.A. to study moths, beyond the Golden Na- 
ture Guide. Shortly following Dover's 1968 reprint, 
Ronald W. Hodges (1971) gave us the first, Sphingi- 
dae, of several volumes in the exquisite Moths of 
America North of Mexico series. Charlie Covell’s ea- 
gerly awaited Field Guide to Eastern Moths in 1984 
was aimed at a general audience. 

My fellow moth-enthusiast acquaintances often 
mention their awe at which the Europeans publish 
smartly produced, high quality, pithy, and easy to use 
books on moths, not only of a general nature, but also 
for the specialist. Such are two new books from Apollo 
Books; The Geometrid Moths of Europe Volume 1, and 
The Sesiidae of Europe. This is a review of the former. 

Wow!! What a book! This is terrific. My first impres- 
sion was “They did it again” or to be more specific 

“Peder Skou did it again.” Skou’s Apollo Books, long a 
publishing house of high quality scientific poole is 
here again with another tome—one that will be refer- 
enced for many years to come. The European users of 
this book will have much more to say about the 
nomenclature and systematics. As a general volume on 
Geometridae, it is terrific. As a general volume on 
moths it is terrific. As a general volume on much of 
Lepidoptera, it is terrific. 

Axel Hausmann, the author of Volume 1 is the Edi- 
tor-in-Chief of a project planned to cover 6 volumes. 
Volume | is full of general information and an appeal 
for assistance with the remaining volumes. 

When I first looked at this hoot I liked it. Then I 
scanned and began to see the goodies. Then I read it 
in detail, and I was overwhelmed. This is a must read 
for all persons interested in Geometridae anywhere. 
The introductory sections are an extremely valuable 
important read for all persons interested in Lepi- 
doptera. 

There are many reasons why I am so enthusiastic: 

1. The distribution maps provide two sets of data si- 
multaneously. Black dots show the exact sites of speci- 
mens examined for this publication. Gray shading 
shows where the species might be seen. The gray 
shading represents extrapolated distributions based on 
ideal habitats, known dispersal patterns, and previ- 
ously published distributions. 

2. The introductory sections provide information on 
many ancillary subjects, of interest to persons studying 


VOLUME 56, NUMBER 1 


Lepidoptera, but most often available only in more 
specialized publications. Do you want to know more 
about “r-strategists,” “K-strategists,” and the “r-K- 
continuum” of “bionomic strategy?” Information can 
be found here. 

Do you want to know more about the dynamics of 
Zoogeography including areal expansions and areal re- 
gressions? Do you want to know more about mor- 
phology (structures) with definitions and illustrations 
of the areole, connate and anastomosis veins, sexual 
and seasonal dichroism, ommatidium, chaetosema, 
flagellum, tergum, sternum, papillae anales, collicu- 
lum, hypognathous, holometabolism, and cremaster 
without the need for several other books? It’s all here. 

3. The concept of cladistic analysis, with its own jar- 


gon (three examples are monophyly, paraphyletic, and | 


plesiomorphic), often appears in journals and at meet- 
ings of Lepidopterists. For the initiated, including 
most persons with an avocational interest in butterflies 
and moths, the strange language is impenetrable for 
understanding. Not any more. Hausmann provides a 
dictionary of many common terms and their meanings 
followed by the application of the concept to the phy- 
logeny of the Geometridae. This is not a course on 
cladistics, but it provides often-used terms in a context 
providing quick overview. These few paragraphs can 
be indispensable for understanding what the authors 
of more complicated papers are saying. 

4, Hausmann explains the concepts of variation, sea- 
sonal variation, sexual variation, normal variation, all of 
which are richly illustrated with color photos. 

5. A combination of photographs and drawing are 
used to illustrate structural features of adults, anten- 
nae, legs, genitalia, larvae, pupae, eggs, and other mor- 
phological characters. We often look at drawings be- 
cause we understand a picture is worth 1000 words. 
Hausmann provides all methods of explaining and in- 
terpreting important features of Lepidoptera and 
Geometridae. 

6. The concepts of species, subspecies, clines, hy- 
brids, and normal variations are discussed. 

7. I really like the sections on Taxonomy and 
Nomenclature. The taxonomy of the subfamilies of 


Geometridae is given followed in a few pages with a 
brief lesson on nomenclature and the Code of Zoolog- 
ical Nomenclature (1999). Hausmann refers to the 
newest version of the code and defines terms such as 
synonymy, homonymy, priority, invalid names, avail- 
ability of names, and the different kinds of types. We 
often see and use these terms. Here they are defined 
for quick reference. The caution that all is not always 
as it seems is also given in a few Problematic cases. 

You can have all this for the low price of $61.00 
USD, but wait, there’s more. The first 79 pages of the 
book are worth the purchase price, but when you buy 
this book you also get 130 pages of very well prepared 
species accounts, eight color plates, 18 pages of geni- 
talia drawings, a systematic checklist, 20 pages of bib- 
liography, and two indices, an Index to subjects and 
taxa, and an Index to Scientific Names. Hausmann 
doesn't miss anything. The text is full of literature cita- 
tions for persons wanting more information. He indi- 
vidually credits each photographer and artist for the 
excellent text illustrations. It’s all here. 

The only thing missing is keys. I like well-prepared 
keys, and this book has no keys. The well-prepared di- 
agnoses and discussions of similar species are a good 
replacement for the keys. 

Overall the volume is pleasingly and expertly con- 
structed. The decorative cloth cover is attractive, and 
the book is Smythe sewn. A Smythe sewn book has 
threads through the folds of the paper on the inside 
margin. A Smythe sewn book lies flat when open, and 
individual pages never fall out. The insides are clean, 
and clearly written with a comfortable typeface and 
large font. The color illustrations are very sharp. The 
production of the book is consistently very high quality. 

I highly recommend this book for all persons inter- 
ested in Lepidoptera. For persons studying Geometri- 
dae the taxonomy presented here is required. For all 
other persons, this is a great example of the qualities of 
a very good book. The information within is very valu- 
able. This book is very good value. 


Eric H. METZLER, The Ohio Lepidopterists, 1241 
Kildale Sq. N., Columbus, Ohio 43229-1306, USA. 


Journal of the Lepidopterists’ Society 
56(1), 2002, 48 


MANUSCRIPT REVIEWERS FOR 2001 (VOLUME 55) 


Manuscript reviewers are anonymous contributors to the scientific rigor, clarity and quality of text and illustrations 
in the papers published by the Journal of the Lepidopterists’ Society. The reviewers’ input is always valuable, but 
their swift response was especially critical during a time when we needed to expedite the publication of volume 55. 
Let us hope such enthusiastic response continues to allow the Journal to grow in readership. On behalf of all au- 
thors and editorial staff of this Journal, respectful acknowledgements are given to the reviewers listed here. Aster- 
isks denote those who reviewed two or more manuscripts. 


Adamsky, David; Washington, DC 
Aiello, Annette; Panama City, Panama 
Austin, George; Las Vegas, NV 
Balogh, George; Portage, MI 
*Bowers, Deane; Boulder, CO 
Brower, Andy; Corvallis, OR 

*Brown, John; Washington, DC 
Brown, Richard; Mississippi State, MS 
Burns, John; Washington, DC 

Chew, Francie; Medford, MA 
Debinski, Diane; Ames, IA 

*DeVries, Phil; Milwaukee, WI 
*Dirig, Robert; Ithaca, NY 

Fahrig, Lenore; Ottawa, Canada 
Hodges, Ronald; Eugene, OR 
Lafontaine, Don; Ottawa, Canada 
Land, Michael; Brighton, England 
Landolt, Peter; Wapato, WA 

McCabe, Timothy; Albany, NY 
Metzler , Eric; Columbus, OH 


Miller, William; Saint Paul, MN 
Nijhout, Fred; Durham, NC 
*Oliver, Jeff; Louisville, CO 
Peigler, Richard; San Antonio, TX 
*Penz, Carla; Milwaukee, WI 
Porter, Adam; Medford, MA 
Powell, Jerry; Berkeley, CA 

Prudic, Katy; Louisville, CO 
*Robbins, Robert; Washington, DC 
Rubinoff, Dan; Berkeley, CA 
Rutowski, Ron; Tempe, AZ 

Solis, M. Alma; Washington, DC 
Sperling, Felix; Edmonton, Canada 
Wagner, David; Storrs, CT 
Wahlberg, Niklas; Stockholm, Sweden 
*Weiss, Martha; Washington, DC 
Weller, Susan; Saint Paul, MN 
White, Raymond; Palo Alto, CA 
Youngstaedt, Elsa; Milwaukee, WI 


EDITORIAL STAFF OF THE JOURNAL 


Carta M. Penz, Editor 
Department of Invertebrate Zoology 
Milwaukee Public Museum 
Milwaukee, Wisconsin 53233 USA 
flea@mpm.edu 


Pur DeVries, Book Review Editor 
Center for Biodiversity Studies 
Milwaukee Public Museum 
_ Milwaukee, Wisconsin 53233 USA 
pjd@mpm-edu 


Associate Editors: 
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Fe.ix Spertinc (Canada), Davin L. Wacner (USA), Curister WikLunp (Sweden) 


sa NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of Lepidoptera study. Categories are Articles, Profiles, General Notes, Techni- 
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Send Journal submissions to the editor at the above address (or electronically to: flea@mpm.edu). Contributors should feel free to 
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Submit manuscripts in triplicate, typewritten, entirely double-spaced, with wide margins, on one side only of white letter-sized pa- 
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Sueprarp, P. M. 1959. Natural selection and heredity. 2nd ed. Hutchinson, London. 209 pp. 

196la. Some contributions to population genetics resulting from the study of the Lepidoptera. Adv. Genet. 10:165-216. 


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Lire ais MATURE STAGES O] 
_. W. Peacock and Jeffrey R. Slotte: 


a eee a 


MOTHS AND BUTTERFLIES 


VoLuME 1, 


© This paper meets the require 


Techie 


} Volume 56 Number 2 
, 30 April 2002 
ISSN 0024-0966 


Journal af the 
Lepidopterists’ Society 


oe Published quarterly by The Lepidopterists’ Society 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


Joun W. Brown, President Susan S. Borkxin, Vice President 
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Cover illustration: Euphaedra edwardsi (Nymphalidae), occurring from Sierra Leone to Nigeria and Cameroon. This specimen was _ 
photographed in Uganda by P. J. DeVries. 


JOURNAL OF 


Toe LeEpiporpTeEeRiIsts’ SOCIETY 


Volume 56 


Journal of the Lepidopterists’ Society 
56(2), 2002, 49-52 


2002 Number 2 


WINSLOW J. HOWARD: PIONEER NEW MEXICO NATURALIST 


STEVEN J. CARY 
202 Solana Drive, Santa Fe, New Mexico 87501, USA 


ABSTRACT. Winslow J. Howard arrived in Santa Fe, New Mexico in 1858. He helped found the original New Mexico Historical Society in 
1859. Over the next 30 years Howard moved between mining towns, operating jewelry, watch-making and assay businesses in several states 
throughout the West. His longest stay in any one place may have been in Silver City, New Mexico, from 1880 to 1887. Howard's natural history 
interests included minerals, fossils, plants and insects. He collected some of the oldest documented butterfly specimens from Colorado and New 
Mexico. Howard was an important early amateur naturalist in the western United States and the first resident butterfly collector in New Mexico. 


Additional key words: _assayer, jeweler, history, naturalist, genealogy, patronym, natural history. 


To modern New Mexico lepidopterists, W. J. 
Howard has been little more than a footnote associ- 
ated with a few late 19th century butterfly records 
from southwestern New Mexico (e.g., Toliver et al. 
1994:401). His role was preserved by Cockerell (1899), 
who discussed several New Mexico butterflies includ- 
ing “14 species collected by Mr. W. J. Howard in Grant 
County, N. M.” Howard's modest lot consisted of Adel- 
pha bredowii Geyer, Eurema nicippe (Cramer), Colias 
eurytheme Boisduval, Zerene cesonia (Stoll), Chlosyne 
lacinia (Geyer), Euptoieta claudia (Cramer), Vanessa 
virginiensis (Drury), Vanessa cardui (Linnaeus), 
Nymphalis antiopa (Linnaeus), Thessalia theona 
(Ménétriés), Thessalia fulvia (W. H. Edwards), 
Danaus plexippus (Linnaeus), and Cercyonis pegala 
(Fabricius). His specimen of T. theona is in the collec- 
tion of the Biology Department at New Mexico State 
University. Most of these taxa are routine in south- 
western New Mexico. Cockerell (1899) offered no 
clues about Howard’s whereabouts, when Howard’s 
specimens were collected, or how they came into 
Cockerell’s possession. This author remained curious 
about Howard's identity and collecting activities in 
frontier New Mexico. 


MATERIALS AND METHODS 


Howard provoked mild curiosity among historians 
and scientists over the years. Rocky Mountain biolo- 
gists (i.e., Brown 1957, 1966) were curious about 
Howard's role as an early collector of specimens, some 


of which became types. New Mexico historians (e.¢., 
Bloom 1943, Torrez 1997) were curious because he 
helped to found the original New Mexico Historical 
Society. Though neither group benefited from the ef- 
forts of the other, their publications represent a collec- 
tion of secondary sources that, when taken together, 
opened doors to valuable primary resources. 

Old newspapers, originally noted by Brown as a key 
resource about Howards activities, also proved vital in 
this present study. Microfiche copies of old newspa- 
pers were examined at the Museum of New Mexico's 
History Library (MNMHL) in Santa Fe, the New 
Mexico Records and Archives Center (NMRAC) in 
Santa Fe, and the Silver City Museum (SCM) in Silver 
City. Minutes of the New Mexico Historical Society 
were an important primary resource cited in Bloom 
(1943) and Torrez (1997). A third major primary 
source was the Special Territorial Census, conducted 
by the U.S. government in 1885 in preparation for 
eventual New Mexico statehood. This was reviewed on 
microfiche at the NMRAC. Information about Howard's 
early professional years in New York could not be lo- 
cated because Tiffany & Co. did not retain employee 
records of that vintage (L. Bann pers. com. 1999). 

Information about Howard from the Special Census 
provided an entry point to genealogical records. Ge- 
nealogical inquiries first were made through various 
internet sites. These led the author to the Reference 
and Genealogy Section of the New Hampshire State 
Library, which led to that state’s Bureau of Vital Statis- 


50 


tics. Detailed research into New Hampshire files was 
not practical from the author's home in New Mexico, 
but that resource warrants future attention. 


RESULTS 


Winslow J. Howard first came to the West in 1858 
(Table 1). His arrival in Santa Fe, New Mexico, is doc- 
umented by the following advertisement, which he 
submitted on 18 December 1858, and which was pub- 
lished in the Santa Fe Weekly Gazette of 5 February 
1859. “Mr. W. J. Howard begs leave most respectfully 
to inform the citizens of Santa Fe and the public in 
general that he has taken a room in the placita of the 
Exchange Hotel for the purpose of repairing fine 
Watches of every description, also Clocks, Music 
Boxes, Jewelry, and other mechanical contrivances. By 
the practical experience of thirteen years in the busi- 
ness, and with the recommendation of the largest Jew- 
elry Establishment in this country—the house of 
Tiffany & Co., New York city—he hopes to merit a 
share of public patronage in his line.” 

Howard's broad interests revealed themselves soon 
thereafter. On 15 December 1859, he was one of 25 
people who met in Santa Fe, the territorial capitol, to 
found the New Mexico Historical Society. Comment- 
ing on the minutes of that meeting, Bloom (1943:242) 
remarked that Howard “was one of the prime movers 
in organizing the Society . . . he himself was made sec- 
retary of the meeting—and it was he who stated the 
object for which they had gathered . .. He was the first 
one named in the charter of incorporation.” On 26 De- 
cember Howard was elected Curator and Librarian 
and was asked to “take charge of and classify all Indian 
antiquities and curiosities; geological and mineralogi- 
cal specimens; geographical maps, and objects of nat- 
ural history” (Torrez 1997). Society records indicate 
that Howard resigned his office on 17 April 1860, and 
left Santa Fe to pursue business interests elsewhere 
(Torrez 1997). 

His destination and activities for the next six years 
were illuminated by Brown (1957:45-47). While re- 
searching pioneer entomologists in Colorado, Brown 
found scattered references to specimens collected by 
Howard. Piecing clues together, Brown deduced that 
Howard was associated with mining camps. By reading 
old newspapers, Brown found an announcement in 
The Western Mountaineer of 19 July 1860, promoting 
Howard’s new business in Denver. By 1862 Howard 
was in Central City, Colorado, where he and a partner 
manufactured jewelry under the name of Howard and 
Colony, whose newspaper advertisements continued 
until January 1865. Brown reported that Howard 
moved back East in late 1865, married, and took up 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


residence in Brooklyn, New York. He returned west in 
October 1866, traveling by stagecoach to Montana via 
Denver. After that, Brown found the trail of his 
“jeweler-naturalist’ more difficult to trace. He re- 
ported evidence of Howard in Prescott, Arizona, in the 
1870's and in Leadville, Colorado, in 1879 (Brown 
1966:127), but he never found Howards trail again. 

Howard's life after 1866 remained a mystery until 
Torrez (1997) placed him in the mining town of Silver 
City, New Mexico, in 1880. Further research by SCM 
staff showed that Howard advertised frequently in lo- 
cal newspapers to promote his assay, jewelry and 
watch-making services (S. Berry, SCM, pers. com. 
1997). He was sufficiently newsworthy to be the sub- 
ject of at least two feature stories. One appeared on 10 
June 1882, in The New Southwest, one of several 
newspapers then operating in Silver City: “[A] collec- 
tion to which we would call special attention is that of 
W. J. Howard, assayer and jeweler, who has spent 
twenty-three years in the Rocky Mountains, and has 
devoted his leisure time to the study of metallurgy and 
natural history and who is a member of several scien- 
tific institutions.” This article detailed his “rare and 
choice selections of crystallizations of metals and min- 
erals.” His was “the largest and finest collection of fos- 
sil shells ever collected in New Mexico from various 
localities, cemented on cards with their generic 
species named, besides butterflies and other insects, 
pressed flowers, leaves, etc.” 

The year 1883 witnessed the arrival in Silver City of 
telephones, electric streetlights and a branch railroad 
from the main line at Deming (Berry & Russell 
1995:24-25). One newspaper article that year featured 
Howard as a prominent collector and dealer of ancient 
coins, which he occasionally sold in New York. Other 
details of Howard's life in Silver City were docu- 
mented in the Special Territorial Census of 1885 
(NMARC archives). A Silver City census-taker 
recorded “Howard, W.J.,” as a married, literate, white 
male whose profession was assayer. The Census found 
Howard living in a boarding house with predominantly 
younger, unmarried men including a miner, a tinsmith, 
a laborer, a merchant, a butcher and three saloonkeep- 
ers. This arrangement was routine in frontier mining 
towns and was consistent with having a permanent 
home and wife in the East. 

The Census listed Howard’s birthplace as New 
Hampshire and the birthplaces of his father and 
mother as Massachusetts and New York, respectively. 
Howard's age was stated as 57 years, indicating he was 
born ca. 1828. This information provided a starting 
point for researching genealogical records. Searches 
for a Winslow J. Howard born in New Hampshire ca. 


VOLUME 56, NUMBER 2 


1828 were conducted on the internet in October 2001, 
but hours of automated searching through these con- 
siderable genealogical resources and databases pro- 
duced no match. The Reference and Genealogy Sec- 
tion of New Hampshire State Library (NMSL) 
searched some of their files at the author's request. Li- 
brary staff found no reference to Winslow Howard in 
the New Hampshire federal census for 1850, in 
Stearns’ “Genealogy of New Hampshire,” or in pub- 
lished Howard genealogies in the Library’s collection 
(Z. Moore pers. com. 23 October 2001). Similarly, the 
New Hampshire Bureau of Vital Statistics (NHBVS) 
found no birth record for Winslow J. Howard 
(NHBVS pers. com. 8 November 2001). 

Failure to locate Winslow J. Howard in New Hamp- 
shire genealogical records reflects two basic obstacles. 
First, genealogical records are incomplete. The 
NHBVS reported that many vital events of that era 
were never recorded (W. R. Bolton pers. com. 8 No- 
vember 2001). Second, “Howard” and “Winslow” were 
popular family names in New England. Internet 
searches revealed one intermarriage between the two 
families, and there likely were others. The NHSL re- 
ported Howard surnames in 34 different New Hamp- 
shire towns in the early 19th century (Z. Moore pers. 
com. 23 October 2001). 

Howards financial fortunes waxed and waned with 
Silver City’s mining-based economy. In 1886, he was 
tax delinquent on property worth $400 (SCM archives 
1998). Economic troubles worsened in 1887, causing 
one of Silver City’s three banks to fail (Berry & Russell 
1995:35). That year, Howard’s regular newspaper no- 
tices assumed a different tone, stating in July 1887 that 
he sold his Silver City business and, in August, that he 
began a professional association with George Williams 
& Co. in Deming. There, Howard’s newspaper trail 
ended. The author tried to trace Howard through the 
Deming Headlight, a newspaper that began publishing 
in 1882, but issues from 1883 to August 1888 could not 
be located in archives. Review of issues from Septem- 
ber 1888 to April 1889 (MNMHL archives) disclosed 
no ads by Howard or by George Williams & Co. 


DISCUSSION 


Despite living on the western frontier, far from most 
professional scientists, Howard did not conduct his 
natural history work in a vacuum. He was a member of 
the Entomological Society of Philadelphia (Brown 
1957) which later became the American Entomologi- 
cal Society. He collected the Colorado specimen used 
to describe a new species of bee (Cresson 1863) and 
he supplied Montana plant specimens to Asa Gray at 
Harvard (Brown 1957). Butterflies collected by 


51 


Howard were among the oldest documented Colorado 
specimens used by Tryon Reakirt to compile the first 
list of Colorado butterflies in 1866 (Brown 1957). 

During Howard's time in New Mexico, he was 
linked to University of Kansas entomologist Francis H. 
Snow. In August 1884, Prof. Snow and his students vis- 
ited Silver City to collect insects (S. J. Cary & R. Hol- 
land in prep.). Reporting on this expedition, Snow 
(1885:65) acknowledged “Mr. W. J. Howard of Silver 
City for many favors.” Snow reported 13 butterflies 
and long lists of moths and beetles, remarking that the 
“electric lights at Silver City were very attractive to in- 
sects” (Snow 1885:69). One particular moth was listed 
as “Daritis sp. a superb species obtained from Mr. W. 
J. Howard, Silver City” (Snow 1885:66). For identi- 
fication of unfamiliar Lepidoptera collected on this 
Bexpedition, Snow (1885:65) relied on Henry Ed- 
wards. In describing the new moth as “Daritis thetis, 
Klug, var. Howardi” (Arctiidae: Pericopinae), Edwards 
(1886[1887]:165) noted that “About 2 years ago, I re- 
ceived from Mr. Wilson [sic] Howard, two specimens 
of this magnificent moth, which had been taken by 
him in New Mexico.” Linkage of the type specimens 
from their collection by Howard, through Snow, to 
Edwards’ description of D. howardi now warrants re- 
striction of its type locality to Silver City, Grant 
County, New Mexico. No other patronyms were 
traced to Winslow J. Howard. 

It has been suggested that Howard may have col- 
lected other specimens that survive today as anecdotal 
frontier New Mexico butterfly reports, for example 
from the Sacramento Mountains. While the timing of 
Howard's presence in New Mexico makes this connec- 
tion theoretically possible, the author is unaware of 
any specific information to support it. Mining was not 
a big part of the frontier economy in the Sacramentos, 
making it is difficult to imagine what would have at- 
tracted Howard to the area or how he could have sup- 
ported himself as an assayer. Similarly, the author is 
not aware of any insect specimens collected by 
Howard during his 1859 sojourn in Santa Fe. Never- 
theless, it is clear that more remains to be learned 
about Howard; additional research may disclose spec- 
imens and relationships that are not apparent today. 


CONCLUSIONS 


In addition to being a professional jeweler and as- 
sayer, Howard was an active naturalist who collected 
specimens that helped scientists describe the flora and 
fauna of the West. Howard appreciated the scientific 
value of his specimens, researched their identities and 
recorded generally where they were collected. Speci- 
mens collected by Howard ca. 1884 are among the 


earliest reliable butterfly reports from southwestern 
New Mexico (Cockerell 1899). He helped University 
of Kansas Prof. F. H. Snow collect insects in Silver City 
in 1884 (Snow 1885). Although genealogical inquiries 
bore no fruit in this study and no evidence was found 
that he fathered any children, Howard’s name lives on 
in the beautiful arctiid moth, Daritis howardi (Ed- 
wards 1886 [1887]). 

The activities, contributions and lives of professional 
scientists of the historical period are documented in 
their publications and through records at their aca- 
demic institutions. In comparison, the lives of most 
amateur naturalists and collectors usually were not sys- 
tematically recorded, even though they supplied much 
of the raw material with which their professional con- 
temporaries worked. It has been challenging to assem- 
ble the story of Winslow J. Howard because his faint 
footprints are scattered across several states, several 
decades and several scientific disciplines. Though his 
origins remain obscure, it is hoped that the above in- 
formation contributes to the record of this important 
naturalist in New Mexico and the West. 


ACKNOWLEDGEMENTS 


The author is grateful to former New Mexico State Historian 
Robert Torrez, who catalyzed this article by providing the clue that 
parted the fog surrounding W. J. Howard in New Mexico. Susan 
Berry and Jim Carlson of the Silver City Museum were very helpful 
in review of Museum archives. Charles Martinez, of the New Mex- 
ico Archives and Records Center, helped identify, retrieve and in- 
terpret historical material. Tomas Jaehn assisted in research of files 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


in the Museum of New Mexico’s History Library. Jane Ruffin looked 
for evidence of Howard at the Library of the Academy of Natural 
Sciences in Philadelphia. Greg Forbes found one extant Howard 
butterfly specimen in the collections of New Mexico State Univer- 
sity. Richard Holland flagged Daritis howardi as a potential W. J. 
Howard patronym and Ron Hodges helped track down its original 
description. Ray Stanford shared his library. Tim Cary and Susan 
Berry critiqued early versions of the manuscript. The author is 
grateful to Richard Holland and an anonymous reviewer for con- 
structive comments. 


LITERATURE CITED 


Berry, S. & S. A. RUSSELL. 1995. Built to last; an architectural his- 
tory of Silver City. Revised 2nd ed. Silver City Museum Society. 
132 pp. 

Bioom, L. B. 1943. Historical Society minutes, 1859-1863. NM 
Hist. Rev. 18(2):247-311. 

Brown, F. M. 1957. Two early entomological collectors in Col- 
orado. Entomological News 68:41-47. 

. 1966. David Bruce (1833-1903) and other entomological 
collectors in Colorado. J. New York Entomol. Soc. 74(3):126-133. 

COCKERELL, T. D. A. 1899. Notes on some New Mexico butterflies. 
Canadian Entomol. 31:65-66. 

CRESSON, E. T. 1863. Proc. Entomol. Soc. Phila. Vol. 2. 

Epwarbs, H. 1886 [1887]. Apparently new forms of N. American 
Heterocera. Entomologica Americana 2(9):165-166. 

SNow, F. H. 1885. Lists of Lepidoptera and Coleoptera collected in 
New Mexico by the Kansas University scientific expeditions of 
1883 and 1884. Trans. Kansas Acad. Sci. [X:65-69. 

TOLIVER, M. E., R. HOLLAND & S. J. Cary. 1994. Distribution of 
butterflies in New Mexico (Lepidoptera: Hesperioidea and Pa- 
pilionoidea). 2nd ed. Publ. by R. Holland. 399 pp. 

TorREZ, R. J. 1997. “Voices from the Past.” Newspaper column in 
Round the Roundhouse. Issue of June 19, 1997.6. 


Received for publication 6 January 2001; revised and accepted 17 
November 2001. 


Journal of the Lepidopterists’ Society 
56(2), 2002, 53-61 


FIVE NEW SPECIES OF DALLA FROM COLOMBIA AND ECUADOR (HESPERIIDAE) 


STEPHEN R. STEINHAUSER 
4715 26th Avenue East, Bradenton, Florida 34208, USA 


ABSTRACT. Five new species of the Heteropterine genus Dalla Mabille, 1904 are described and their male (and female where available) 
genitalia illustrated. Three of these species, Dalla disconnexa, new species, D. vista, new species, and D. celsus, new species aré from 
Ecuador, the other two, D. wardi, new species and D. pedro, new species, from northeastern Colombia. 


Additional key words: _ genitalia, transtilla, harpe, Heteropterini, cypselus group, caenides group. 


When Evans compiled his catalog of the Hesperi- 
inae of America (Evans 1955), there were in the 
BMNH collection 1949 Dalla specimens representing 
71 of the then known 75 species and 41 subspecies. Of 
these, only 29 specimens were females from 15 differ- 
ent species, and two of these females are the unique 
types of their species. He divided these 75 species into 
seven species groups, based on wing and leg charac- 
ters, but not genitalia, resulting in some very unrealis- 
tic associations. Since the publication of Evans’ mas- 
terpiece, 20 new species and two new subspecies have 
been described (D. pota and D. cola Bell, 1959; D. 
ramirezi Freeman, 1969; D. roeveri, Miller & Miller, 
1972; D. nubes, D. pincha, D. xantha, D. bos, D. pura, 
D. simplicis, D. puracensis, D. puracensis quindio, D. 
puracensis cotopa and D. calima Steinhauser, 1991a; 
D. kemneri Steinhauser, 1991b; D. steinhauseri Free- 
man, 1991; D. freemani, Warren, 1997 and the five de- 
scribed below). Two subspecies, D. epiphanaeus supe- 
rior Draudt, 1923 and D. lalage lethaea (Schaus, 1913) 
have been raised to specific rank by Steinhauser 
(1991a:5, 13), and one species, D. dividuum (Dyar, 
1913) raised from synonymy with D. ligilla (Hewitson, 
1877) by Freeman (1968:61) bringing the totals to 98 
species and 41 subspecies. 

In human-disturbed sites, males of the genus Dalla 
frequently gather on vertical or near vertical wet con- 
crete surfaces, especially recently set concrete to feed 
on the mineral rich moisture exuding therefrom. They 
also congregate on wet soils where cattle have gath- 
ered, feeding on the nitrogen rich urine. I have found 
that urine added to vertical steep faces is a good bait, 
but I have never found females at any of these sites. 
They are either very rare or keep themselves well hid- 
den, probably busy laying eggs to raise the next gener- 
ation of males. Dalla species are not often found below 
1500 m elevation, and usually above 2500 m. I know 
nothing of Dalla life history, food plants or immature 
stages, but it is probable that the larvae may feed on 
various grasses as recorded for Carterocephalus palae- 
mon (Pallas, 1771) by Tietz (1972:501) and Scott 
(1986:425), and for Piruna pirus (Edwards, 1878) and 
P. aea (Dyar, 1912) by Opler (1999:415-416). 


While curating Hesperiid material at the Allyn Mu- 
seum of Entomology, comparative genitalic examina- 
tion with superficially similar taxa in conjunction with 
previous studies indicated that several of the many 
unidentified hesperiids specimens were indeed new 
species. As stated by Judith E. Winston (1999:115), 
“Once you have . . . satisfied yourself that the organism 
you are studying does indeed represent an unde- 
scribed species, your aim is publication. Only if it is 
named and described acceptably in a scientific publi- 
cation will the species name be available for you and 
others to use. Descriptions of new species are still an 
important part of publication in the field of taxonomy.” 
Therefore, in an effort to better document the Neo- 
tropical lepidopteran fauna, five new species in the 
genus Dalla Mabille, 1904, are described below. Nine 
other new species and two new subspecies in the 
genus were described earlier (Steinhauser 1991a, b). 


DALLA MABILLE, 1904 


Diagnosis. The genus Dalla is one of the six het- 
eropterine genera occuring in the New World. Most of 
its 98 species (this includes the five new ones herein 
described) are essentially montane in habitat and cen- 
tered primarily in the Andes of South America, but ex- 
tending also into Central America and Mexico. Evans 
(1955), the most recent reviser of the genus, used ten 
characters, unfortunately none of them genitalic, to 
distinguish among the six genera of the New World 
Heteropterini, which he included in the Hesperiinae 
as the Carterocephalus group. He distinguished Dalla 
from the other five (Carterocephalus, Piruna, Darda- 
rina, Butleria and Argopteron) by the apiculus of the 
antennae being “gradual, sharply pointed” rather than 
“blunt, more or less flattened and compressed at tip” 
as in the other five (Evans 1955:9-10). The other nine 
characters were used in various combinations to dis- 
tinguish among the other five genera. None of their 
various states pertains uniquely to Dalla, which shares 
a relatively long antenna (equal or greater than half the 
forewing costa) with Argopteron, Butleria and Darda- 
rina; spined midtibiae shared with Carterocephalus, 


54 


Piruna, Dardarina and Butleria; relatively short palpi 
(equal to head, rather than longer) shared with Darda- 
rina; antennal nudum usually of more than 11 seg- 
ments rather than less, shared with Butleria; nudum 
longer than half the antennal club, shared with Dar- 
darina and Butleria; antennal club not grooved, shared 
with all but Argopteron. I found the other three char- 
acters used by Evans to apply inconsistently. For fur- 
ther comments on Evans’ classification see Stein- 
hauser (1991b:40-42). 


MATERIALS AND METHODS 


I have followed Evans’ arrangement of the genus 
into groups in the placement of these new species, de- 
spite its unreality. Wing measurements are given to the 
nearest 0.5 millimeter, since I find it impossible to de- 
termine the exact wing base position on a mounted 
specimen more closely. Genitalic dissection tech- 
niques and terminology are the same as used by Stein- 
hauser (1989). Wing venation follows the system of 
Miller (1972). The male genitalia of all five and the fe- 
male genitalia of the one with a known (or probable) 
female are illustrated. Two of these new species are 
from the Santa Marta region of northeastern Colom- 
bia. The other three, two of which I had previously 
misidentified as Dalla connexa Draudt, 1923, are from 
Ecuador. All of these specimens are deposited in the 
Allyn Museum of Entomology. 


Dalla wardi Steinhauser, new species 
(Figs. 1, 2, 15) 


Male. Head: Blackish brown above; palpi hairy, grizzled black 
and white, third segment (missing in holotype) slender, porrect, 
black, nearly hidden in hairs of second segment. Antennae about 
half costa, shaft prominently checkered black and yellow, club black, 
yellow at base, nudum brown, 11 segments in holotype (right an- 
tenna glued to paper triangle on pin), 12 segments in paratype; ter- 
minal segment short, pointed. Thorax: Blackish brown above, ful- 
vous beneath. Thoracic appendages: Legs fulvous; foretibiae with 
long slender, brown epiphyses reaching and slightly overlapping 
tarsi; mid and hindtibiae spined, midtibiae with single pair of spurs, 
hindtibiae with two, the upper smaller. Wings: Dorsal surface: 
Forewing dark blackish brown, a few scattered yellowish hairs in 
basal third; yellowish white hyaline spots as follows: three contigu- 
ous, subequal subapical spots separated by dark veins in R,-R,, 
R,-R, (smallest) and R-M, (largest), their inner edges in an arc 
cone ex proximad; triangular lower cell spot nearly reaching radius, 
centered between origins of R, and R,; large nora anel spot in 
Cu,-Cu,, its edges in line with those of the cell spot and separated 
Gan it only by the dark cubital vein; small (circular in the holotype, 
rhomboid in the paratype) spot in mid M,—Cu,, somewhat larger 
than the largest subapical spot. These hyaline spots are bordered by 
a very narrow line of orange yellow scales. In addition to the hyaline 
spots, there is an opaque rhomboid-to-near-triangular yellow spot in 
mid Cu2—2A, not reaching Cu, and separated by about half its width 
from the spot in Cu,—Cu,, its outer edge about in line with the inner 
edge of the eon’ cell plus Cu,—Cu, spot. Fringes concolorous, 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


shading to paler brown and somewhat ochreous at torus. Hindwing 
same dark brown as forewing, with a few orange yellow hairs in the 
basal quarter, and bearing a prominent, sharply defined, somewhat 
ovoid central orange spot, ee by dark veins, in the cell, ex- 
treme base of Rs—M,, M,— ,-M,,, extreme base of M,—Cu,, 
base of Cu,—Cu, and Cu ee fae Sah of the spot in Cu, Cis Hal 
Cu,-1A is shifted slightly basad from the rest of the 5p Fringes 
ochreous to orange, slightly paler at tornus. Ventral surface: 
Forewing centrally blackish brown, costa and apex broadly reddish 
brown, more or less heavily scaled yellowish in distal half of costal 
cell. Hyaline spots as above; opaque spot in Cu2—2A much larger 
than above, clear pale yellow, shares entire caudal edge of spot in 
Cu,-Cu,, concave distally, convex proximally. Fringes reddish 
brown, shading to greyish at tomus. Hindwing reddish brown, dark 
grey in 2A—3A and anal cell. Central spot as above, clear orange yel- 
low, sharply defined. Additional rather obscure opaque yellow spots 
in Sc+R,—Rs about one third way from base, and subtornally in 
Cu2-2A. Fringes concolorous at apex, shading to ochreous at tornus. 
Abdomen: Blackish brown above, fulvous beneath. Genitalia: Very 
similar to D. mesoxantha (Pl6tz, 1884), D. xantha Steinhauser, 1991, 
D. merida Evans, 1955 and D. frater (Mabille, [1879]). Tegumen 
slender, oval, not hollowed above; uncus rather short, subequal to 
tegumen, slender and narrowly and shallowly bifurcate in dorsal 
view; in lateral view, not projecting dorsad at juncture with tegumen, 
slightly hooked at distal end, bearing prominent dorsal hair tuft. 
Gnathos well sclerotized, smooth, extends distad to about mid uncus. 
Valvae symmetrical, 2.4 times longer than wide, 1.8 times length of 
combined tegumen and uncus; harpe projects prominently cephalad 
as a slender dentate process with a straight rather than concave dor- 
sal edge, completely overlapping the obliquely uptummed distal por- 
tion of the ampulla which does not extend dorsad beyond harpe and 
bears a slender, inwardly projecting curved flange at its base. Penis 
slender, slightly shorter than valvae, distally broadened and dentate 
on left side; phallobase extremely short; single small, doubly dentate 
cornutus. Saccus very short, triangular; juxta and transtilla prominent. 

Female. Unknown. 

Wing measurements. Male forewing 13 x 7 mm (paratype) to 
13.5 x 7.5 mm (holotype). 

Type material. Holotype ¢, Colombia: Magdalena; § km E of 

San Pedro, 2550 m, 13-XII-1973, P. Ward & A. Forsyth, bearing the 
following labels: printed and hand printed white label, COLOM- 
BIA: 8 km E of San Pedro Dept. Magdalena 10°55’N, 73°58’W 2550 
m. 13.X1I.1973 P. Ward, A. Forsyth; hand printed yellow label, [P.S. 
Ward photo slide No.] 7-19; white paper triangle with right antenna 
glued thereupon; printed and hand printed white label, Allyn Mu- 
seum Acc. 1999-9; printed and hand printed red label, HOLOTYPE 
3 Dalla wardi S.R. Steinhauser; printed and hand printed white la- 
bel Allyn Museum Photo No. 990724-13,14. There is one male 
paratype, same data as holotype; both of which are deposited in the 
Allyn Museum of Entomology. 

Etymology. This handsome skipper is named for its discoverer, 
Dr. Philip S. Ward. 

Discussion. Dalla wardi keys out to D. mesoxantha 
in Evans’ (1955) key to the species of Dalla. It is a 
member of Evans’ cypselus group and is most closely 
related to D. mesoxantha, D. xantha, D. frater and D. 
merida, but is smaller: 13-13.5 mm for D.wardi, 
14-15 mm for D. frater, 15-15.5 mm for D. mesoxan- 
tha and D. xantha. The ventral hindwing reddish 
brown ground color of D. wardi is like that of D. 


merida and similar to that of D. frater and D. meso- 


VOLUME 56, NUMBER 2 


D. wardi & HT 


D. disconnexa & PT 


ee 


D. vista & HT 


59 


D. disconnexa 6 HT 


D. disconnexa 2 PT 


D. pedro d HT 


UL 
1 cm 


D. celsus & HT 


Fics. 1-14. New Dalla species (odd numbers dorsal, even numbers ventral): 1, 2—Dalla wardi, new species, 3 holotype; 3, 4—Dalla dis- 
connexa, new species, 3 holotype; 5, 6—Dalla disconnexa, new species, ¢ paratype; 7, 8—Dalla disconnexa, new species, 2 paratype; 9, 10— 
Dalla vista, new species, 3 holotype; 11, 12—Dalla pedro, new species, 3 holotype; 13, 14—Dalla celsus, new species, 3 holotype. 


xantha, but unlike the dark brown of D. xantha. There 
is some question about the identity of D. mesoxantha 
(see Steinhauser 1991a:10), but until proven other- 
wise, I use the BMNH specimen from the Kaden col- 
lection marked “Type” as its model, though I have not 
seen its genitalia. The ventral hindwing maculation of 
D. wardi is like that of D. mesoxantha and D. merida, 
but the color of the central spot, both ventral and dor- 
sal is more orange in D. wardi than in D. mesoxantha 
but not as deep orange as in D. merida. There is a 
more or less complete row of postdiscal pale spots in 
D. frater, lacking in D. wardi. 

In the male genitalia, the uncus of D. wardi, viewed 
dorsally, is considerably more slender than in D. 
mesoxantha and D. xantha, but not as slender as D. 
frater; I have not seen the genitalia of D. merida, and 


Evans’ sketch does not make this feature clear. The 
forward edge of the uncus in D. wardi and D. frater 
does not project over the tegumen as it does in D. 
mesoxantha and D. xantha. The penis of D. wardi, like 
that of D. frater, is more slender than in D. xantha and 
D. mesoxantha, but, like them, much shorter relative 
to the valva length than D. frater (0.89 times valva for 
D. wardi and D. mesoxantha, 0.93 times for D. xantha, 
but 1.13 times for D. frater; the penis length of D. 
merida is not known). The dentate dorsal process of 
the harpe is very slender and slightly concave dorsally 
in D. mesoxantha, somewhat thicker in D. xantha, D. 
frater and D. wardi, strongly concave in D. frater, very 
strongly concave in D. merida according to Evans’ 
sketch, only slightly concave in D. xantha, but straight 
in D. wardi. Only in D. wardi is the ampulla com- 


56 


pletely overlapped by the harpe, it projects dorsad be- 
yond the harpe in the others. 


Dalla disconnexa Steinhauser, new species 
(Figs. 3-8, 16, 17) 


Male. Head: Blackish brown; palpi missing in type series of four 
males. Antennae (broken in holotype) slightly longer than half costa, 
shaft prominently checkered black and yellow, club black above, yel- 
low beneath, nudum dark brown, 12 segments in two male 
paratypes with complete antenna, terminal segment long, pointed. 
Thorax: Blackish brown above, fulvous beneath. Thoracic ap- 
pendages: Legs pale fulvous; foretibiae with minute brown central 
epiphyses; mid and hindtibiae spined, midtibiae with single pair of 
spurs, hindtibiae with two. Wings: Dorsal surface: Forewing dark 
blackish brown, a few scattered yellowish hairs in basal third; 
opaque yellow-orange (holotype) to pale yellowish white spots as fol- 
lows: three contiguous, subequal subapical spots not separated by 
dark veins in R,—R,, R,-R, and R,-M.,, their outer edges in a line di- 
rected toward ail ener a more or ees triangular lower cell spot, 
centered approximately between origins of R, and R,; large rhom- 
boid spot in Cu,—Cu,, its inner edge in line with that of ey cell spot 
and separated froin it only by the dark cubital vein; small, more or 
less quadrate spot in M,—Cu,, subequal to the subapical spots. These 
spots are slightly paler yellow-orange in one paratype and pale yel- 
lowish white in the other two. Fringes basally concolorous, distally 
paler brown, still paler at tomus. Hindwing same dark brown as 
forewing, with a few orange yellow (holotype) hairs in the basal 
quarter (pale yellow or missing in the paratypes), and bearing a 
prominent, fairly well defined, elongate ovoid central yellow-orange 
spot, undivided by dark veins and narrowed at its proximal end 
which nearly reaches the wing base, in the cell, M,—M,, M,—M,, ex- 
treme base of M,-Cu,, base of Cu,—Cu, and Cu, <A aie porto of 
the spot in Cu, Cnn, and Cu,-1A is shifted slightly basad from from 
the rest of the spot leaving a small dark notch distally on its rear 
edge. As on the forewing, the spot color varies in the paratypes. 
Fringes pale brown, shading to more orangish at tornus (yellowish to 
whitish in the paratypes). Ventral surface: Forewing centrally 
blackish brown, slightly paler in anal cell; costal cell, costa, apex be- 
yond subapical spots and termen to Cu, broadly reddish brown. 
Opaque spots as above, slightly paler; spot in Cu,—Cu, extended 
patchily to 2A or 1A, variable. Fringes reddish brown. Hindwing 
reddish brown, dark grey with sprinkling of pale yellowish scales in 
1A-2A, 2A-3A and anal cell. Central spot as above, but extending 
into Sc+R,-Rs, filling the basal three fourths of that space, which 
bears, slightly distad of midpoint, a vague, reddish brown spot over- 
scaled yellow. There is some postdiscal, scattered yellow scaling, but 
not organized into a spot band. Fringes concolorous, slightly paler at 
tornus. Abdomen: Blackish brown above, fulvous to whitish be- 
neath. Genitalia: Illustrated by Hayward (1943) and Evans (1955) 
as D. connexa (Hayward and Evans, nec Draudt 1923: see Mielke 
1993:620, figs. 67-71 for correct rendition of D. connexa genitalia). 
Tegumen in dorsal view, oval, centrally constricted, not hollowed 
above; uncus rather short, in dorsal view oval, elongated distally to a 
narrow, bluntly pointed nose, in lateral view projecting dorsad and 
cephalad over tegumen, distally slightly hooked, bearing a dense 
dorsal hair tuft. Gnathos well sclerotized, smooth, exeniting caudad 
to beyond mid uncus, where it is surmounted dorso-distally by a 
broad, distally bifurcate, shagreened process wider than uncus and 
projecting caudad subequally. Valvae symmetrical, three times as 
long as wide, 1.7 times as long as combined tegumen/uncus length; 
harpe projects dorso-distally as a slender, curved process, coarsely 
dentate terminally and along its forward edge, only slightly overlap- 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


ping the ampulla which is evenly rounded and upturmed dorsad, and 
distally elongated more or less evenly with the harpe. Penis slender, 
long (1.2 to 1.3 times valva length); phallobase short; cornutus a 
single, small, dentate process. Juxta and transtilla very prominent; 
juxta narrowly pointed cephalad; transtilla developed into two long 
pointed arms projecting caudad. 

Female. Head: As male; palpi missing from both paratypes, an- 
tennae of the one paratype with complete antennae, as male. Tho- 
rax and thoracic appendages: Thorax and legs as male. Wings: 
Generaily as male, dorsal surface wing markings slightly paler; ven- 
tral surface ground color, especially the hindwing, much paler, cen- 
tral spot pale yellow, postdiscal area densely covered with pale yel- 
low scales, leaving a narrow reddish brown distal border to the 
central spot, most prominent in M,—M, and M,—M., and a very nar- 
row reddish brown terminal border before the faage Genitalia: 
Lamella postvaginalis a narrow, distally concave seleroured process. 
Lamella antevaginalis consists of two lateral lobes. Antrum a moder- 
ately sclerotized, rather long, oval tube extending well forward of the 
lamella antevaginalis; ductus bursae with a slender, rather obscure 
internal sclerotized process at about mid point, well forward of the 
antrum. Corpus bursae consists of two spherical sacs in tandem con- 
nected by a narrow membranous neck; the forward sac simple, the 
more caudad sac with long, slender internal spines forming lateral 
signa. Ductus seminalis connected to mid ductus bursae ventrally. 

Wing measurements. Male forewing 11 x 5.5 to 12 x 6.5 mm 
(holotype); female forewing 11.5 x 6 to 12 x 6 mm. 

Type material. Holotype ¢, Ecuador: Cotopaxi; Milimbanco 
3900 m, ix.1971, R. de Lafebre, bearing the following labels: printed 
white label, ECUADOR: COTOPAXI Milimbanco, 3900 m; ix.1971 
R. de Lafebre; printed white label, A.C. Allyn Acc. 1972-2; printed 
and hand printed white label, Genit. Vial SRS-2543; printed and 
hand printed red label, HOLOTYPE ¢ Dalla disconnexa S.R. Stein- 
hauser; printed and hand printed white label, Allyn Museum Photo 
No. 990724-7,8. There are three male and two female paratypes all 
by the same collector: 1 d Ecuador: Pichincha; Niebli, NW slope of 
Vol. Pichincha, 3500 m, viii-1971; 1 ¢ Ecuador; Pichincha; Vol. 
Antisana, 2950 m, vii-1971; 1 ¢ Ecuador: El Oro; Bellavista, 550 m, 
v-1971; 1 2, Ecuador: Tungurahua; Bellavista nr. Banos, 1900 m, xii- 
1970; 1°, Ecuador: Tungurahua, San Antonio, 1950 m, ix-1971. 

Etymology. I have named this insect D. disconnexa because of 
its great similarity to, but differences from, D. connexa. 


Discussion. See discussion of the new species de- 
scribed immediately below. 


Dalla vista Steinhauser, new species 
(Figs. 9, 10, 18) 


Male. Head: Blackish brown above; palpi hairy, basally black, 
grizzled black and white distally, third segment (missing in holotype 
and one male paratype) slender, porrect, black, nearly hidden in 
hairs of second segment. Antennae slightly longer than half costa, 
shaft prominently checkered black and yellow, club black above, yel- 
low beneath, nudum dark brown, 12 segments in only male paratype 
with complete antenna (tips of apiculi broken off on holotype), ter- 
minal segment long, pointed. Thorax: Blackish brown above, ful- 
vous beneath. Thoracic appendages: Legs pale fulvous; foretibiae 
with minute brown central epiphyses; mid and hindtibiae spined, 
midtibiae with single pair of spurs, hindtibiae with two. Wings: 
Dorsal surface: Forewing dark blackish brown, a few scattered yel- 
lowish hairs in basal third; opaque yellow-orange spots as follows: 
three contiguous, subequal subapical spots not separated by dark 
veins in R,-R,, R,-R, and R.—M,, their outer edges in a line directed 


VOLUME 56, NUMBER 2 57 


aS 17 


disconnexa 
-——4 


Fics. 15-20. Dalla species genitalia (scale line = 1 mm). 15—Dalla wardi, 3 paratype (Genit. Vial SRS-4804): a) tegumen, uncus, vincu- 
lum and associated structures, lateral; b) tegumen and uncus, dorsal; c) tegumen, uncus and gnathos, ventral; d) saccus, ventral; e) right valva, 
interior lateral; f) penis, cormutus, transtilla and juxta, dorsal; g) penis, transtilla and juxta, lateral; 16—Dalla disconnexa, 3 holotype (Genit. Vial. 
SRS-2543): a) tegumen, uncus, vinculum and associated structures, lateral; b) tegumen, uncus and gnathos, dorsal; ¢) tegumen, uncus and 
gnathos, ventral; d) saccus, ventral; e) right valva, interior lateral; £) penis, cornutus, transtilla and juxta, dorsal; g) penis, cornutus, transtilla and 
juxta, lateral; 17—Dalla disconnexa, ? paratype (Genit. Vial SRS-5281), ventral. 18—Dalla vista, 3 holotype (Genit. Vial SRS-5282): a) tegu- 
men, uncus, vinculum and associated structures, lateral; b) tegumen and uncus, dorsal; e) tegumen, uncus and gnathos, ventral; d) saccus, ven- 
tral; e) right valva, interior lateral; f) penis, comutus, transtilla and juxta, dorsal; g) penis, cornutus, transtilla and juxta, lateral; 19—Dalla pe- 
dro, 3 paratype (Genit. Vial SRS-4805): a) tegumen, uncus, vinculum and associated structures, lateral; b) tegumen, uncus and gnathos, dorsal; 
c) tegumen, uncus and gnathos, ventral; d) saccus, ventral; e) right valva, interior lateral; f) penis, comutus, transtilla and juxta, dorsal; g) penis, 
cornutus, transtilla and juxta, lateral. 20—Dalla celsus, 3 holotype (Genit. Vial SRS-4797): a) tegumen, uncus, vinculum and associated struc- 
tures, lateral; b) tegumen, uncus and gnathos, ventral; ec) tezumen, uncus and gnathos dorsal; d) saccus, ventral; e) right valva, interior lateral; 
f) penis, cormutus and juxta, dorsal (drawn from memory). 


58 


toward mid termen; a more or less triangular lower cell spot, cen- 
tered approximately between origins of R, and R,; large rhomboid 
spot in Cu,—Cu,, its inner edge in line with that of the cell spot and 
separated from it only by the dark cubital vein; small spot in 
M,—Cu,, subequal to the subapical spots (rhomboid in the holotype 
and one paratype and not reaching the base of M.—Cu,, triangular in 
one paratype and reaching the base). Fringes light greyish brown 
with an orange tinge toward the tornus. Hindwing same dark brown 
as forewing, with a few orange-yellow hairs in the basal quarter, and 
bearing a prominent, fairly well defined, elongate ovoid central yellow- 
orange spot, undivided by dark veins and narrowed at its proximal 
end which nearly reaches the wing base, in the cell, M|—M,, M.—M,, 
extreme base of M,—Cu,, base of Cu,—Cu, and Cu,—1A; the portion 
of the spot in Cu,—Cu, and Cu,-1A is shifted slightly basad from 
from the rest of the spot leaving a small dark notch distally on its 
rear edge. Fringes ochreous to orange, shading to more orange at 
tornus. Ventral surface: Forewing centrally blackish brown, 
slightly paler in anal cell; costal cell, costa, apex beyond subapical 
spots and termen to Cu, broadly reddish brown. Opaque yellow- 
orange spots as above, slightly paler; subapical spots narrowly bor- 
dered distally by slightly darker reddish brown; spot in Cu,—Cu, ex- 
tended broadly to 2A, yellow-orange to vestigial vein 1A, whitish in 
1A-2A. Fringes reddish brown. Hindwing reddish brown, dark grey 
with sprinkling of pale yellowish scales in 1A-2A, 2A-3A and anal 
cell. Central spot as above, more or less overscaled reddish brown, 
poorly defined, marked with a vague reddish brown line at cell end. 
There is a very vague yellowish postdiscal spot-band from about 
Rs—M, to Cu,—Cu,, which may be reduced to a few yellow scales. 
Fringes concolorous, slightly paler at tornus. Abdomen: Blackish 
brown above, fulvous beneath. Genitalia: Tegumen slender, oval, 
not hollowed above; uncus rather short, in dorsal view oval, elon- 
gated distally to a narrow, bluntly pointed nose; in lateral view pro- 
jecting dorsad and cephalad over tegumen, distally slightly hooked, 
bearing a dense dorsal hair tuft. Gnathos well sclerotized, smooth, 
extending caudad to about mid uncus, where it is surmounted dorso- 
distally by a rounded, distally somewhat excavate, shagreened 
process narrower than uncus. Valvae symmetrical, three times as 
long as wide, 1.6 times as long as combined tegumen/uncus length; 
harpe projects dorso-distally as a slender, smooth pointed process 
with a single, centrally placed, short, inwardly projecting tooth; am- 
pulla distally elongated, evenly rounded, bearing a narrow, curved, 
inwardly projecting flange at its base, overlapped by harpe process 
which does not reach dorsad beyond mid ampulla. Penis slender, 
long (1.4 times valva length); phallobase short; cornutus a single, 
small, dentate process. Juxta and transtilla very prominent; juxta nar- 
rowly and bluntly pointed cephalad; transtilla developed into two 
long pointed arms projecting caudad. 

Female. Unknown. 

Wing measurements. Male forewing 11.5 x 6 mm (one 
paratype) to 12 x 6 mm (holotype and one paratype). 

Type material. Holotype ¢, Ecuador: El Oro; Bellavista 550 m, 
V-1971, R. de Lafebre, bearing the following labels: printed white 
label, ECUADOR: EL ORO; Bellavista, 550 m; v.1971 R. de 
Lafebre; printed white label, A.C. Allyn Acc. No. 1972-6; printed 
and hand printed red label, HOLOTYPE ¢ Dalla vista S.R. Stein- 
hauser; printed white label, Genit. Vial No. SRS-5282; printed and 
hand printed white label, Allyn Museum Photo No. 990724-3,4. 
There are two male paratypes, all same collector as holotype, 1 d 
same data as holotype, 1 d Ecuador, Tungurahua; Bafios, 1850 m, xii- 
1970. The holotype and paratypes are deposited in the Allyn Mu- 
seum of Entomology. 

Etymology. The name vista is based on the last part of the name 
of the village where the holotype was collected. 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Discussion. I had determined a series of seven 
males (two dissected) and two females (one dissected) 
from Ecuador at the Allyn Museum as D. connexa, 
based on Evans’ (1955) key and description and his 
and Hayward’s (1943) genitalia drawings. When later 
compared with Mielke’s (1993) illustration of the gen- 
italia of the lectotype of D. connexa, it appeared that 
those two males whose genitalia I had examined might 
be different; they had a prominent, bifurcate transtilla 
like that illustrated by both Evans and Hayward, and 
the dorsal harpe process was prominently and coarsely 
dentate along its entire forward edge, unlike that 
shown by Mielke with only some fine terminal denta- 
tion. Mielke did not illustrate the juxta and transtilla, 
and did not remember seeing (pers. com. 1999) a 
prominent, bifurcate transtilla. Upon dissecting the 
abdomens of the remaining specimens in the series, I 
discovered that three of the males were entirely differ- 
ent (D. vista); the other two were the same as the two 
I had originally determined as D. connexa, newly de- 
scribed above as D. disconnexa. I am treating the fe- 
males as D. disconnexa because, although the central 
yellow spot of the hindwing beneath is more or less 
obscurely defined as in the males of D. vista, and not 
as well defined as in D. disconnexa males, it does ex- 
tend fully into Sc+R,—Rs, and does not have a reddish 
brown bar at cell end. It is possible, however, that they 
may belong to D. vista instead. 

It is very probable that the skippers determined as 
D. connexa by both Hayward and Evans, at least those 
whose genitalia were examined, are D. disconnexa 
rather than D. connexa. I have before me, however, a 
photograph of an undissected BMNH specimen from 
Colombia, determined by Evans as D. connexa, that I 
believe is probably D. vista. 

Superficially, I find it very difficult to distinguish 
among the three species. All are about the same size 
(11-12 mm forewing) and have essentially the same 
wing markings. Comparing the Seitz illustration of D. 
connexa (Vol. 5: pl. 179e), and a photo taken by Mielke 
of the lectotype, with the series before me of D. vista 
and D. disconnexa, and a photograph of the BMNH 
specimen mentioned above, determined by Evans as 
D. connexa, I find the following slight and perhaps in- 
consistent differences: 

Dorsal surface: spot in Cu,—Cu, of forewing may be 
slightly larger in D. vista and D. disconnexa than in D. 
connexa. Color of the spots in D. disconnexa varies 
from yellow-orange to yellowish white; D. connexa is 
illustrated as yellow-orange similar to D. vista. The 
hindwing central spot of D. connexa and D. disconnexa 
is slightly longer (extends closer to termen) than that 
of D. vista. 


VOLUME 56, NUMBER 2 


Ventral surface: in the Seitz illustration, the hind- 
wing central spot of D. connexa, which may be of the 
female, is very large, nearly white and fairly well de- 
fined. That of D. disconnexa, which may be whitish, is 
equally well defined, but very much smaller, whereas 
that of D. vista is rather poorly defined and generally 
slightly darker. The central spot in D. vista does not ex- 
tend into Sc+R,-Rs as in D. disconnexa and has a red- 
dish brown bar at cell end, missing in D. disconnexa. 

Both of these new species belong in Evans’ (1955) 
caenides group, and both will key to D. connexa in his 
key to the Dalla species. 


Dalla pedro Steinhauser, new species 
(Figs. 11, 12, 19) 


Male. Head: Dark brown above; palpi missing from both holo- 
type and paratype; antennae reach to slightly beyond mid costa, 
shaft prominently checkered yellow and black, club black above pale 
yellow beneath, nudum brown, 12 segments, terminal segment 
short, rounded. Thorax: Dark brown above, beneath clothed in 
long pale fulvous hairs. Thoracic appendages: Legs clothed in 
long pale fulvous hairs; foretibiae with very small contra epiphyses, 
“il and hindtibiae spined, midtibiae sath single pair of spurs, 
hindtibiae with two pairs. Wings: Dorsal surface: Forewing dark 
brown with scattered ochreous hair scales in basal third. Small 
opaque, subapical yellow-orange spots in R,—R,, R,-R, and R.—M, in 
a line directed toward mid termen; slightly larger (up to 1 mm), 
widely separated, round, opaque yellow-orange spots in M,—Cu, and 
Cu,—Cu,. Fringes concolorous, shading to paler dull orange at tor- 
nus. Hindwing same dark brown as forewing, with scattered ochre- 
ous hair scales in basal third; small central orange spot in cell end ex- 
tending into basal Rs-M,, M,-M, and M,—M.,. Fringes dull orange. 
Ventral surface: Fonecting ne danke brown ae above, slightly paler 
in anal cell, broadly rufous at costa, apex and termen to Cu,. Opaque 
yellow-orange spots as above, those in M,—Cu, and CnC, much 
larger, quadrate; an additional opaque pale yellow spot in Cu2—2A 
adjoining the spot in Cu,—Cu,. Fringes as above. Hindwing rufous, 
black brown in anal cell; faint darker brown postdiscal spot band 
from Sc+R,—Rs to Cu,—Cu,. Fringe rufous with a faint dark brown 
hairline at its base. Abdouien: Dark brown above, rufous beneath. 
Genitalia: Tegumen broad in dorsal view, somewhat quadrate in 
lateral view, slightly hollowed dorsally. Uncus slender, entire, slightly 
hooked at distal end, projects over tegumen where it bears a very 
dense hair tuft. Gnathos broad, sclerotized, smooth, but bearing a 
large, bifurcate, dentate dorsal process that extends caudad as far as 
does the uncus. Valvae symmetrical, long (1.5 times combined tegu- 
men/uncus length), narrow (length 2.5 times its greatest width); 
harpe projects dorsad as a short dentate nose, reaching mid ampulla, 
which is large, evenly rounded and extending caudad nearly as far as 
harpe. Penis long, slender, curved to left, terminally widened to left; 
phallobase very short; cornutus a small, monodentate plate. Juxta 
and transtilla prominent, transtilla with short, bilateral caudally di- 
rected projections. Saccus a small slender triangle. 

Female. Unknown. 

Wing measurements. Male forewing 12.5 x 6.5 mm (holotype) 
to 13 x 7 mm (paratype). 

Type material. Holotype 3, Colombia: Magdalena; 8 km E of 
San Pedro, 2550 m, 13-XII-1973, P. Ward & A. Forsyth, bearing the 
following labels: printed and hand printed white label, COLOM- 
BIA: 8 km E of San Pedro Dept. Magdalena 10°55’N, 73°58’W 2550 


m. 13.XII.1973 P. Ward; hand printed yellow label [P.S. Ward photo 
slide No.] 7-21; printed and hand printed white label, Allyn Mu- 
seum Acc. 1999-9; printed and hand printed red label, HOLOTYPE 
3 Dalla pedro S.R. Steinhauser; printed and hand printed white la- 
bel Allyn Museum Photo No. 990724-11,12. There is one male 
paratype, Colombia: Magdalena; 10 km E of San Pedro, 2900 m, 23- 
XII-1973, P. Ward; both types are deposited in the Allyn Museum of 
Entomology. 
Etymology. This skipper is named for its locale, San Pedro. 


Discussion. Dalla pedro does not fit well into 
Evans’ 1955 key to the species of Dalla. I place it ten- 
tatively in Evans’ caenides group on the basis of its 
rather peculiar genitalia, similar to D. connexa Draudt, 
1923, D. ticidas (Mabille, 1897), D. caenides (Hewit- 
son, 1868), D. bos Steinhauser, 1991, D. mora Evans, 
1955 and D. carnis Evans, 1955, none of which evenly 
faintly resembles D. pedro superficially. Dalla pedro 
bears some superficial similarity to several of the more 
sparsely marked Dalla species, but is immediately rec- 
ognized by its distinctive genitalia. 


Dalla celsus Steinhauser, new species 
(Figs. 13, 14, 20) 


Male: Head: Black-brown above; palpi and antennae missing. 
Thorax: Black-brown above, reddish brown beneath. Thoracic Ap- 
pendages: Legs dark brown, clothed in ochreous hairs and scales; 
fore and mid legs missing, hindtibiae spined with two pairs of spurs. 
Wings: Dorsal surface: Forewing black-brown with scattered 
ochreous scaling in the basal quarter, most prominent along the 
costa. Opaque yellow-orange spots subapically in R.—R, (smaller), 
R,-R, and R.—M,, in a line directed toward mid termen; broad cell 
spot across mid cell, its somewhat excavate outer edge centered be- 
tween R, and R,; large, somewhat elongate spot in Cu,—Cu,, its 
outer edge convex, its inner half conjoined to the cell spot. Fringe 
paler brown. Hindwing same black-brown as forewing, overlain with 
orange hairs in basal one third, bearing a large, tear-drop-shaped or- 
ange spot nearly reaching wing base in cell, bases of Rs-M,, M,—M,, 
M,-M,, M,-Cu,, Cu,—Cu, and extending slightly into Cu iA a 
Sonik —Rs. ences Berman orange. V ‘oral eee Forewing cen- 
trally dull black, reddish brown along costa to about cell end ne at 
apex; spots as above, some yellow scaling at costa above cell spot; 
rather faint narrow yellowish spot in Cu2—2A, adjoining spot above 
it in Cu,—Cu,, narrow at Cu,, broader at 2A. Fringe red-brown, paler 
basally. Eitecofing reddish brown, blackish in Cu2—2A and anal cell: 
spot from above allows very faint, indistinct. Fringe concolorous, 
shading to orange at tomus. Abdomen: Blackish rows above, red- 
dish brown beneath. Genitalia: Tegumen short, broad, globular; 
uncus short, broad, dorsally hollowed, its distal end broad, rounded, 
slightly concave centrally and extending laterally as two blunt points; 
the uncus is surmounted by two large circular lobes, densely hairy 
and projecting prominently dorsad. Gnathos shagreened, broad as 
uncus, rounded, slightly excavate centrally at caudal end, and ex- 
tends caudally beyond uncus. Valvae symmetrical, harpe distally an 
upright process which is finely serrate along its distal margin, the 
teeth pointing inward. The harpe extends dorsad subequally with 
the rather squared, dorsally pointed ampulla, which, in interior view, 
overlaps the harpe rather than the more usual harpe overlapping the 
ampulla. Penis (lost while transferring to second watch glass and 
drawn immediately from memory) long, slender, with very short 
phallobase, slightly flared to the left distally and with a single small, 


60 


dentate cornutus. Juxta (lost with penis and drawn from memory) 
prominent, projecting prominently cephalad; transtilla (lost with pe- 
nis and drawn from memory), not projecting prominently caudad. 

Female. Unknown. 

Wing measurements. Holotype male forewing 15 x 7.5 mm. 

Type material. Holotype 3, Ecuador: Chimborazo; Atzata- 
pungu, 4100 m, vi, 1976, R. de Lafebre, bearing the following labels: 
printed white label, ECUADOR: CHIMBORAZO Atzatapungu, 
4100 m vi.1976; R. de Lafebre; printed white label, A.C. Allyn Acc. 
1976-8; printed and hand printed white label, Genit. Vial SRS-4797; 
printed and hand printed red label, HOLOTYPE 4 Dalla celsus S.R. 
Steinhauser; printed and hand printed white label, Allyn Museum 
Photo No. 990724-1,2. Known only from the holotype which is de- 
posited in the Allyn Museum of Entomology. 

Etymology. This skipper is named celsus, Latin for lofty, due to 
the high altitude of its type locality. 


Discussion. In Evans’ (1955) key to the species of 
Dalla, D. celsus falls into the caenides group, between 
D. seirocastnia Draudt, 1923 and D. pantha Evans, 
1955, its hindwing spot extending slightly into 
Sc+R,—-Rs, unlike D. seirocastnia, but not reaching 
Sc+R,, as in D. pantha. Another superficially similar 
species is D. simplicis Steinhauser, 1991 which lacks 
the ventral surface forewing spot in Cu2—2A. Genitali- 
cally, D. celsus is closest to D. seirocastnia, D. pura- 
censis Steinhauser, 1991 and D. ochrolimbata Draudt, 
1923 in having prominent dorsal lobes surmounting 
the uncus. Dalla ochrolimbata was placed by Evans in 
the quadristriga group because of its nearly complete 
lack of dorsal surface markings. Dalla celsus differs 
from both D. seirocastnia and D. ochrolimbata in the 
terminal shape of the uncus: broad and rounded in D. 
celsus and D. puracensis; with a bluntly pointed nose 
in D. seirocastnia and D. ochrolimbata. It also differs 
from them in the general shape of the valvae, which 
are terminally deeply excavate in D. seirocastnia and 
D. ochrolimbata. There are many other differences, 
but these serve to separate the species. 


CONCLUSIONS 


The genus Dalla, in fact the entire Heteropterini 
tribe, is in need of a phylogenetic revision. For ex- 
ample, I suspect that a few species currently in Dalla 
may belong in Piruna. It is also possible that some 
named Dalla taxa may be mere infraspecific variants; 
Evans (1955:20), speaking of his agathocles group, 
notes “The genitalia of the following 8 species are too 
alike to confirm their validity as species.” Two of those 
eight species were then given an additional two sub- 
species and one of them, an additional three, making 
15 taxa with essentially identical genitalia. Whereas 
this is entirely possible, it does seem like a ripe field 
for more detailed investigation. Unfortunately, at the 
present time, lack of comparative material from varied 
locations plus the extreme scarcity of females in col- 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


lections, makes such a study very difficult. If there are 
more undescribed species hiding in collections, they 
should be described and published to add to the data 
available for a possible revision. Whether this will re- 
sult in subdividing Dalla into more than a single genus 
or to the establishment of sub-genera, I cannot say. 
There is a good possibility of arranging the species into 
groups based on genitalic similarity, shape and form of 
the antennal apiculus and club, form of the foretibial 
epiphyses and very probably other characters as well, 
certainly female morphology and immature biology. I 
have accumulated a fair amount of data, drawings and 
photos, which I will gladly share with anyone who de- 
cides to tackle this project. 


ACKNOWLEDGMENTS 


Thanks are due and given to Olaf H. H. Mielke, Universidade Fed- 
eral do Parana, Curitiba, Brasil for supplying a photograph of the lec- 
totype of Dalla connexa and for discussion via e-mail about Dalla gen- 
italia; to Philip S. Ward for donating the type series of D. wardi and D. 
pedro to the Allyn Museum; to Jacqueline Y. and Lee D. Miller for 
their continued support and critical and helpful review of the second 
draft of this paper; to C. Don MacNeill for his review of the first draft 
of the manuscript; to Carla Penz and an anonymous reviewer for help- 
ful suggestions that I have tried to use to improve the content, and to 
my very patient wife, Josie, for her patience and forbearance. 


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61 


taxa in the neotropical Hesperiidae. Part I, Pyrginae. Bull. Allyn 

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Received for publication 20 February 2001; revised and accepted 
4 December 2001. 


Journal of the Lepidopterists’ Society 
56(2), 2002, 62-65 


THE COLLAPSED EGGS FOUND IN THE BURSA COPULATRIX OF A PLUM MOTH, ILLIBERIS 
ROTUNDATA JORDAN (ZYGAENIDAE: PROCRIDINAE): AN UNUSUAL EGG RESORPTION SYSTEM? 


CHIHARU KOSHIO, YOSHIHISA TANAKA AND SHIN-ICHI KUDO 


Naruto University of Education, Takashima, Naruto 772-8502, Japan 


ABSTRACT. Ina plum moth, Illiberis rotundata Jordan (Zygaenidae), collapsed eggs and empty chorions were usually found in the bursa 
copulatrix. Effects of the number of these eggs on female longevity were analyzed with female body weight, body weight of her mate, and fe- 
male fecundity. The results of multiple regression analysis showed that females with more eggs in the bursa copulatrix lived longer. It is most 
likely that eggs are resorbed in the bursa copulatrix and used for the survival of the females. This moth seems have an unusual egg resorption 


system. 


Additional key words: oocyte, longevity. 


Egg resorption, a specific type of reproductive tactic 
in which oocytes degenerate instead of being laid as 
eggs, has been reported in many insects (Bell & Bohm 
1975), including lepidopteran species from groups 
such as Heliconiinae (Dunlap-Pianka et al. 1977) and 
other Nymphalidae (Boggs & Ross 1993). 

Illiberis rotundata Jordan (Zygaenidae) is a univol- 
tine and diurnal moth. Larval host plants include cher- 
ries, plums and other Rosaceae trees. Adults emerge 
from May to June in western Japan, and copulate and 
oviposit repeatedly almost for a month, taking only wa- 
ter (CK unpublished data). During the process of a 
study on mating behavior of I. rotundata, we found 
many collapsed eggs in the bursa copulatrix of females. 
In this paper, we describe this unusual phenomenon 
and suggest a possible function of these eggs. 


MATERIALS AND METHODS 


We collected pupae of Illiberis rotundata in Naruto 
City (34°11’N, 134°35’E) in May 2000. Each indi- 
vidual was kept separately in a paper cup (7 cm diam- 
eter, 7 cm depth) placed in a constant condition room 
at 21 + 1°C with a 15L:9D photoperiod. After emer- 
gence, all adults were weighed using an electric bal- 
ance (Sartorius AG) with an accuracy of 0.01 mg. 
Nineteen females were allowed to copulate once one 
day after emergence and five females were kept un- 
mated through their lives (the laboratory females). 
Each mated female was moved to and kept in a plastic 
case (9 cm diameter, 5 cm depth) supplied with a 
single fresh cherry leaf (Prunus x yedoensis) and wa- 
ter. These females were allowed to lay eggs until 
death. Each cherry leaf was renewed every evening af- 
ter the number of eggs laid by the female on that day 
had been checked. We dissected these females soon 
after their deaths and examined the contents of the 
bursa copulatrix. In 2001, we also collected and dis- 
sected females from the field (the wild females). 

In order to reveal the function of the eggs in the 
bursa copulatrix, we examined their effects on female 
longevity. We selected four factors that might affect fe- 


male longevity: female body weight (body weight of 
each female when she emerged), male body weight 
(body weight of her mate when he emerged), fecun- 
dity (total number of eggs each female laid during her 
lifetime) and the number of eggs (in any condition, see 
results) in the bursa copulatrix. Before further analy- 
ses, each value of female body weight, male body 
weight, fecundity and the number of eggs (+1) in the 
bursa copulatrix was log-transformed. 

We analyzed correlation structures between these 
factors and longevity. Multiple regression analysis was 
also used to estimate only the direct effects of each 
factor on female longevity, using female longevity as 
the dependent variable, and the four factors as the in- 
dependent variables. Data were analyzed using the 
StatView 5.0 (SAS Institute Inc.). 


RESULTS 


We dissected 22 laboratory females and 21 of them 
(95.5%) had eggs in their bursa copulatrix (Fig. 1). We 
found a few spheroid shaped eggs, several collapsed 
eggs, and many empty eggs including fragmented 
chorions (Fig. 1b). The spheroid eggs were observed 
near the ductus bursa, whereas the empty eggs were at 
the bottom of the bursa. In one female, a spheroid egg 
was also observed in the ductus seminalis. We could 
not recognize any distinct spermatophore. The fat 
body had almost been depleted at death. 

The number of eggs, including spheroid, collapsed 
and empty eggs, in the bursa copulatrix varied from 
zero to 52 in the laboratory females (Mean + SD = 
11.4 + 12.8, N = 19). In many cases, the bursa con- 
tained some other small fragmented chorions, thus the 
counted number of eggs seems to be underestimated. 
Among those 19 females, longevity was 23.6 + 6.0 days 
(Mean + SD), fecundity was 461.1 + 177.3, body 
weight was 53.7 + 8.8 mg, and body weight of their 
mates was 31.6 + 3.4 mg. 

The number of eggs in the bursa copulatrix and fe- 
male body weight were positively correlated with fe- 
male longevity (Table 1, Fig. 2), but fecundity was neg- 


VOLUME 56, NUMBER 2 


63 


Fic. 1. The bursa copulatrix containing eggs in the laboratory females. (a) Many eggs contained are visible through the translucent wall of 
the bursa (BC, bursa copulatrix; OV, oviduct; SP, spermatheca). After dissection of this bursa, 9 collapsed and 18 empty eggs were counted. (b) 


Spheroid, collapsed or empty eggs from the bursa of another female. 


atively correlated with female longevity. Fecundity was 
also positively correlated with female body weight (r = 
0.46, P = 0.046). Fecundity and number of eggs in the 
bursa, however, showed no significant correlation (r = 
—0.36, P = 0.13). 

The total multiple regression model was highly sig- 
nificant (R? = 0.82, F = 15.5, P < 0.0001; Table 1). 
Both the number of eggs in the bursa and female body 
weight positively influenced female longevity. Fecun- 
dity negatively affected female longevity, suggesting a 
phenotypic cost of reproduction (see Reznick 1985). 
Male body weight had no effect on female longevity. 

There is no significant correlation between fecun- 
dity and the number of eggs in the bursa, partialling 
out longevity (partial r = 0.129, P = 0.59, N = 20). 

Nine out of 16 wild females that had been collected 
in the field also had collapsed eggs or chorions in their 
bursa copulatrix. Seven of the nine females had one or 
two eggs, while two females had highly degenerated 
chorions that could not be counted. These observa- 
tions indicate that retention of eggs in the bursa copu- 
latrix is not caused artificially by keeping females in the 
laboratory for long periods. However, the number of 


retained eggs found in wild females (Mean + SD = 0.64 
+ 0.75, N = 14) was smaller than that of the laboratory 


females (Mann-Whitney’s U-test, U = 13.5, P < 0.0001). 


DISCUSSION 


Resorption usually occurs in immature eggs (oocytes) 
within the ovarioles (Bell & Bohm 1975). On the other 
hand, the resorption of mature eggs (chorionated 
eggs) has been reported in Heliconiine butterflies 
(Dunlap-Pianka et al. 1977) and other insects (see Bell 
& Bohm 1975). 

In most laboratory females of I. rotundata, col- 
lapsed eggs with chorions were found in their bursa 
copulatrix at death. To the best of our knowledge, such 
a phenomenon has never been documented in Lepi- 
doptera. At this point we might ask, what is the func- 
tion of these eggs? Eberhard (2000) found a mature 
egg or a larva just hatched from the egg in the bursa 
copulatrix in some female Microsepsis armillata 
(Diptera: Sepsidae) flies, and he also reported the 
same phenomenon in other flies. He suggested that 
the egg or larva would prevent intromission by a male, 
even though in these species females are immune to 


TaBLE 1. Results of correlation and multiple regression analysis for female longevity and factors potentially affecting the longevity. The total 
multiple regression model was highly significant (see text). r: correlation coefficient. B: standardized partial correlation coefficient. 


Correlation 
Factor r P 
Female body weight 0.29 0.232 
Male body weight 0.21 0.391 
Female fecundity —0.52 0.023 


Number of eggs in bursa 0.76 


<0.001* 


Multiple regression 


B t Pp 
O48 3.55 0.003* 
0.06 0.53 0.607 

-0.55 -3.68 0.003* 
0.50 3.81 0.002% 


* Significant after using the sequential Bonferroni correction. 


Female longevity (in days) 


0 10 20 30 40 50 60 


Number of eggs in bursa copulatrix 


Fic. 2, The relationship between the number of eggs in the 
bursa copulatrix and female longevity. 


rape. In I. rotundata, females are usually polyandrous 
and these eggs in the bursa copulatrix would not pre- 
vent intromission by males. 

The result of multiple regression analysis shows that 
females which had more eggs in their bursa copulatrix 
lived longer. It is most likely that females consume 
eggs in the bursa copulatrix and use them to survive, 
that is, they re-allocate resources from reproduction to 
survival. There are two pieces of indirect evidence to 
support our hypothesis for I. rotundata. First, most of 
the eggs in the bursa copulatrix were highly degener- 
ated. We could see many empty eggs or fragmented 
chorions at the bottom of the bursa, whereas some 
spheroid shaped eggs were observed near the ductus 
bursa. These suggest that egg contents were digested 
and resorbed during retention in the bursa copulatrix. 
The fact that a spheroid egg was in the ductus semi- 
nalis of a female suggests that eggs were transferred 
from the ovary to the bursa copulatrix via the ductus 
seminalis. Second, it is reported that many lepi- 
dopteran females can consume male spermatophores 
in their bursa copulatrix as nutrition for their eggs 
and/or themselves (Boggs & Gilbert 1979, Boggs 
1981). In I. rotundata, no distinct spermatophores 
were detected in the bursa copulatrix of the laboratory 
females. It is possible that not only spermatophores but 
chorionated eggs are also degenerated in the bursa. 

Bell and Bohm (1975) listed many factors promot- 
ing oosorption. For example, the restriction of the 
ovipositional site sometimes increases resorption of 
eggs. In our experiment, each female was restrained in 
a small plastic case with a leaf of the host plant. These 
unnatural conditions might inhibit their oviposition, al- 
though they laid eggs readily on the leaf or the wall of 
the case. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


In some insects oocytes are quickly resorbed in the 
absence of mating. Illiberis rotundata females usually 
mate repeatedly under field conditions (CK unpub- 
lished data). In our experiment, however, females 
were allowed to mate only once. When females are 
prevented from mating, they may consume some of 
their eggs in order to live longer and possibly achieve 
additional matings. Through multiple matings, these 
females might gain additional nutritional and/or ge- 
netic benefits (Arnqvist & Nilsson 2000, Jennions & 
Petrie 2000). 

Resorption can occur in Lepidoptera in response to 
qualitative or quantitative nutrient deficiencies (Dun- 
lap-Pianka et al. 1977, Boggs & Ross 1993). Adults of 
I. rotundata take only water, and thus no additional 
nutrients from food are available for survival. Never- 
theless they can live for relatively long periods, some- 
times more than three weeks. Egg resorption in this 
species seems to be an effective system for obtaining 
additional nutrients at the expense of reproduction. 

Wild females also had collapsed eggs or degener- 
ated chorions in their bursa copulatrix, but the number 
of these eggs and chorions was very low. Two reasons 
for this are to be considered: first, the age of the wild 
females is uncertain and they may have been dissected 
at a younger age than the laboratory females, which 
were dissected after living out their lives. If there is a 
positive correlation between the number of eggs in 
bursa copulatrix and female age, younger wild females 
would have less eggs in the bursa than older laboratory 
females. Second, wild females may have copulated re- 
peatedly until the point of collection, while the labora- 
tory females mated only once. If males provide nutri- 
tional investment to females during copulation, mating 
frequency will have an effect on egg resorption by fe- 
males. 

Why, then, do I. rotundata females resorb eggs in 
their bursa copulatrix not in ovarioles like other in- 
sects? One possible reason could be to eliminate re- 
mains after egg resorption. If resorption of chorion- 
ated eggs occurs in ovarioles, the remains of resorbed 
eggs, such as chorions, should be eliminated to facili- 
tate the passage of the following eggs. Some wasps 
oviposit empty chorions to solve this problem (see Bell 
& Bohm 1975). Disposal of the remains in the bursa 
copulatrix seems to be less costly than in ovarioles, be- 
cause remains can be left in the bursa copulatrix. But 
it is still unknown why they resorb chorionated eggs. 


ACKNOWLEDGMENTS 


We thank A. R. Chittenden for his suggestions improving an ear- 
lier draft of this paper. We also thank C. L. Boggs for her constructive 
comments and C. M. Penz for her editorial help on the manuscript. 


VOLUME 56, NUMBER 2 


LITERATURE CITED 


ARNQVIST, G. & T. NiLsson. 2000. The evolution of polyandry: 
multiple mating and female fitness in insects. Anim. Behav. 
60:145-164. 

BELL, W. J. & M. K. Boum. 1975. Oosorption in insects. Biol. Rev. 
50:373-396. 

Boccs, C. L. 1981. Nutritional and life-history determinations of 
resource allocation in holometabolous insects. Am. Nat. 
117:692-709. 

Boces, C. L. & L. E. GILBERT. 1979. Male contribution to egg pro- 
duction in butterflies: evidence for transfer of nutrients at mat- 
ing. Science 206:83-84. 

Boccs, C. L. & C. L. Ross. 1993. The effect of adult food limita- 
tion on life history traits in Speyeria mormonia (Lepidoptera: 
Nymphalidae). Ecology 74:433-441. 


65 


DUNLAP-PIANKA, H., C. L. Boccs & L. E. GILBERT. 1977. Ovarian 
dynamics in Heliconiine butterflies: programmed senescence 
versus eternal youth. Science 197:487—490. 

EBERHARD, W. G. 2000. Behavior and reproductive status of Mi- 
crosepsis armillata (Diptera: Sepsidae) flies away from oviposi- 
tion sites. Ann. Entomol. Soc. Am. 93:966-971. 

JENNIONS, M. D. & M. Petriz. 2000. Why do females mate multi- 
ply? A review of the genetic benefits. Biol. Rev. 75:21-64. 
REZNICK, D. 1985. Cost of reproduction: an evaluation of the em- 

pirical evidence. Oikos 44:257—-267. 


Received for publication 2 July 2001; revised and accepted 3 De- 
cember 2001. 


Journal of the Lepidopterists’ Society 
56(2), 2002, 66-89 


STUDIES IN THE GENUS HYLEPHILA BILLBERG, I. THE BOULLETI SPECIES GROUP 
(HESPERIIDAE: HESPERIINAE) 


C. DON MACNEILL 
Department of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118, USA 


ABSTRACT. This is the second of three papers on the genus Hylephila Billberg (1820). The first paper introduced the genus and, em- 
phasizing male and female genitalia, defined four species groups, two of which, the ignorans and the venusta groups, were treated in detail. The 
present treatment keys adults; illustrates adults, stigma pockets (male), and g genitalia ( (male and female), and maps the known distribution of the 
species of the boulleti group, seven of which are described as new. Although specimens (especially females) are woefullly scant, their characters 
seem to suggest two subgroups with five known species each. Subgroup I has Hl herrerai , new species, H. pseudoherrerai, new species, H. 
pallisteri, new species, H. blancasi, new species, and H. tentativa, new species. Subgroup II comprises H. shapiroi, new species, H. 
galera Evans, H. boulleti (Mabille), H. rossi, new species, and H. peruana Draudt. Examination of the male and female genitalia is generally 
necessary to be certain of the identity of a specimen, but wing patterns and some structural characters may permit a “good guess” once famil- 
iarity with these skippers is established. Ova of two species were obtained by dissection, and these are described briefly for the first time. 


Additional key words: South America, oreal (of high mountain), altiplano, Andes, dashed Phulia pattern, genitalia (male and female), 


stigma pockets. 


In a previous paper (MacNeill & Herrera 1999) on 
the genus Hylephila Billberg (1820), we introduced 
the genus, emphasized the importance of male and fe- 
male genitalia, and reviewed earlier attempts to illus- 
trate these. We recognized, defined, and provided a 
superficial key to four species groups. We mapped 
three of these groups which are principally Andean, 
and treated two of them, the ignorans group and the 
venusta group, in detail. We alse discussed and figured 
Linka lina (Plétz 1883) owing to confusion about its 
relationship to some high altitude species of Hylephila, 
especially the boulleti group (Shapiro 1994:44, 45), 
which is treated here in detail. Techniques, methods, 
and materials—as well as terminology—used in that 
study also apply in the present paper. A third paper 
(MacNeill in prep.) will treat the remainder of the 
genus, the phyleus group. 

Specimens were examined from several institutions 
listed below, abbreviations are as follows throughout 
the text and figure legends: 

AME: Allyn Museum of Entomology, Florida Mu- 
seum of Natural History, Sarasota, Florida. 

AMNH: The American Museum of Natural History, 
New York, New York. 

BMNH : The Natural History Museum, London, 
England. 

CAS: California Academy of Sciences, San Fran- 
cisco, California. 

CMNH: Carnegie Museum of Natural History, 
Pittsburgh, Pennsylvania. 

IEUM: Instituto de Entomologia, Universidad Met- 
ropolitania de Ciencias de la Educacion, Santiago, 
Chile. 

IML: Instituto de Zoologia, Fundacién Miguel 
Lillo, San Miguel de Tucuméan, Argentina. 

HNHM: Hungarian Natural History Museum, Bu- 
dapest, Hungary. 


LACM: The Natural History Museum of Los Ange- 
les County, Los Angeles, California. 

MUSM: Museo Nacional de Historia Natural, Uni- 
versidad Mayor de San Marcos, Lima, Perit. 

UCD: Bohart Museum of Entomology, University 
of California, Davis, California. 

UFPC: Departamento de Zoologia, Universidade 
Federal do Parana, Curitiba, Brasil. 

USNM: National Museum of Natural History, 
Smithsonian Institution, Washington, D.C.. 


THE BOULLETI GROUP 


The ten species of the boulleti group all look very 
much alike superficially. It is usually necessary to dis- 
sect the genitalia to identify a specimen. These are 
small to medium-sized skippers (forewing length 8.5 
mm-—15 mm). The body is blackish, but the black tegu- 
lae are contrastingly and broadly edged with pale yel- 
low or white. On both surfaces of the secondaries 
(Figs. 32-57) there is a well-defined ray in the cell (at 
least distally) and through spaces M1—M3. Bold black 
spots are characteristic on the ventral surface basally, 
postmedially, and marginally, where they appear to be 
defined or cut by white or pale veins. This gives the 
group an appearance when at rest that oo 
(1985:8—10) has called “the dashed Phulia pattern” — 
common pattern at rest shared by some high eee 
pierids and hesperiids of oreal bogs and meadows, and 
this may well be, as Shapiro suggested, the result of 
convergence for crypsis at rest. He noted that these 
swift flying animals are remarkably difficult to find at 
rest, even when very abundant. 

The species of this group occur in the high Andes 
from central and southern Peri, western Bolivia, and 
north-eastern Chile into northwestern Argentina and 
the pre-Andean Nevadas del Aconquija (the Sierras 
Pampeanas), just west of San Miguel de Tucuman, Ar- 


VOLUME 56, NUMBER 2 


Fic. 1. Diagrams of stigma pockets i in males of three species of 
Hylephila. a, H. pallisteri, new species holotype, PERU, Cuzco, Ol- 
lantaitambo, III-24-47, 9200 ft., C. J. Pallister ( (genitalic dissection # 
é 3811-JH), in AMNH. b, H. pallisteri PERU, Cuzco, Abra Acjan- 
nacu, 3600 m, V-17-84, G. Lamas (genitalic dissection # 6 6227- 
CDM), in MUSM. e, H. blancasi, new species holotype, Santia, aa 
nated 1902, P. forte [sic] (genitalic dissection # d 3873-]H), 
AMNH. d, H. tentativa, new species holotype, PERU, Aiketuen 
Rio Apacheta, 4200 m, 13°21’S, 74°39’W, 1-24-99, G. Lamas, (geni- 
talic dissection # 6 6308-CDM), in MUSM. 


gentina. All of the species in this group except one are 
known from Pert, and the exception almost certainly 
occurs there too. 

Because they are found at high elevations where 
both the weather and access are often limiting, and be- 
cause of their crypsis (see above), these species of 
Hylephila are not often collected and they remain rare 
in collections. I have seen more than five specimens of 
only two of the ten species and females of only five of 
these (three are single specimens), and I have seen 
only one male specimen of four of the ten species. 

Evans (1955) treated three species of this group as 
two subspecies of H. boulleti, (Mabille 1906), H. b. 
boulleti and H. b. peruana Draudt (1923), and one 
new species, H. galera Evans. Seven additional species 
are described in this paper. Given the general paucity 
of material, I must admit to some hesitation regarding 
the possibly presumptive classification presented here 
for this group of Hylephila. 

Based upon the male and female genitalia, the 
species of this group sort rather well into two sub- 
groups. The males of the five species in the first sub- 
group tend to have the anterior half of the uncus 
rather rounded in dorsal view and the gnathos is mas- 
sive and caudoventrally somewhat divergent from the 
uncus in lateral view (Figs. 12-18); two solitary female 
specimens lack a detectable sclerotized eighth sternite 
(Figs. 26, 27). The other subgroup of five species has 
males with the uncus more or less triangular in dorsal 
view, with minute lateral serrations caudally, the 
gnathos not as divergent ventrally from the uncus 
(Figs. 19-25), females have a distinctly sclerotized 
eighth sternite. 


67 


A 


Cc d 


—_ Vi 
oe 
ae BS 

Fic. 2. Diagrams of stigma pockets in males of two species of 
Hylephila. a, H. shapiroi, new species holotype, PERU, Junin, vic. 
Abra Anticona, 4843 m, X-19-83, A. M. Shapiro (genitalic dissection 
# 3 5005-JH), in CAS. b, H. galera holotype, PERU, Junin, Galera 
Pass, 4800 m, II-?-00, (Siang), in BMNH. e¢, H. galera? PERU, 
Junin, Tarma, 3000 m, I-29-72, (genitalic dhesection 2 6 6121-CDM), 
in LACM. d, H. galera? PERU, Yauli, Corpacancha, 4300 m, 


11°29’S, 76°13’W, ie 18-97, R. Acero (genitalic dissection # ¢ 6307- 
CDM), in MUSM. 


ARTIFICIAL KEY TO SPECIES 


l= Malle satdnoriit A SAME ooo occ vacccasedeacanvcescnn0de 2 
1’- Male with a stigma Res Se rach Shad Merve, Shue Peasant bal 3 
2- Hindwing above with fulvous of space M1—M3 a short, broad 
ray, not broadly entering discal cell; below, vannal fuscous 
area wide, broadly entering space Cul-2A ............... 
dd S OOS Sera ron Cea ta eine H. herrerai, new species 
2’- Hindwing above with fulvous of space M1—M3 a long ray 
broadly entering discal cell nearly to base; below, coaarmell 
fuscous area narrow, confined to anal cell and space 2A-3A 
ee iene atime sy ns ear H. pseudoherrerai, new species 
3- Male stigma with yellow microandroconial mass conspicuous 
without magnification, contrasting with other stigmal 
Clementsuee eee rere ce wate asin eee. H. peruana (Draudt) 
3’- Male stigma with microandroconial mass inconspicuous with- 
out magnification, appearing black to tan and not contrasting 


Yntln Qdner caieAmeall EWAN 4 64566005060000000000500008 4 
4- Male with upper element of stigma pocket reduced (Figs. 2a, 
4’- Male with upper element of stigma pocket well developed 

(OST US) eie-tases 04:6. gc. o-ontt.o 6 bd.dln goles oo onc ao .cloen at 6 


5- Very small (FW 8.5 mm). Forewing rather narrow, with up- 
per element of stigma pocket extending distad from vein Cu2 
but not nearly reaching origin of vein Cul (Fig. 2a); hind 
tibia with single pair of spurs ..... H. shapiroi, new species 

5’- Medium sized (FW 13 mm). Forewing broad, with upper el- 
ement of stigma pocket extending distad from a point well 
above vein Cu2 and nearly or quite reaching vein Cul 
(Fig 2b); hind tibia with two pairs of spurs... H. galera Evans 

6- Male forewing with microandroconial mass ae stigma under 
magnification appearing black ...H. pallisteri, new species 

6’- Male forewing with microandr odontal mass of stigma under 


magnification appearing gray, tan, or yellowish ............ if 
7- Forewing above with broad fuscous border deeply cut by 

broad Elkan along veins nearly or quite (O) WEIAONEAM «sp a0 8 
7’- Forewing above with fuscous border not deeply cut to 

termen by fulvous veins, or if deeply cut, narrowly so ..... 10 


8- Forewing above with pale fulvous postmedian band with in- 
ner edge nearly or quite continuously in line with that of 


subterminal spots. ............. H. blancasi, new species 
8’- Forewing above with inner edge of fulvous postmedian 

band offset at vein M3 from sulbiomnniiael fulvous spots ..... fs) 
9- Larger insect (FW 13 mm); forewing broad, not produced; 

{REINO DAUD go og sed cooagecunDvesuacnas H. galera Evans 


9’- Slightly smaller insect (FW 12 mm); forewing somewhat nar- 
row, slightly produced; fulvous warm, allnnos OWING so ccc: 
Be ee aa CEI re H. tentativa, new species 


68 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


LAG AANA 


a b Cc d 
3 


Fic. 3. Diagrams of stigma pockets in males of three species of 
Hylephila. a, H. boulleti, CHILE, Antofagasta, Tatio, 4360 m, J. 
Herrera, in CAS. b, H. boulleti, ARGENTINA, Tucuman, Las Ani- 
mas, Portozuelo de las Animas, 4540 m, I-26-79, (genitalic dissec- 
tion # J 5003-JH), in IML. e, H. rossi, new species holotype, PERU, 
Puno, 10 mi. N. Ayaviri, HI-1-51, E. S.Ross & A. E. Michelbacher 
(genitalic dissection # ¢ 3821-JH), in CAS. d, H. peruana, PERU, 
Junin, Pachachaca, 4000 m, V-20-79, G. Lamas, (genitalic dissection 
#¢d 3871-JH), in MUSM. 


10- Hindwing above with broad fulvous postmedian macular 
band extending at least into space Rs—M1, fulvous pale to 
very pale yellowish ................-. H. boulleti (Mabille) 

10’- Hindwing above with narrow fulvous postmedian macular 
band not extending beyond spaces M1—M3, fulvous dark, 
nearly OHMS poccesoscosogowooses H. rossi, new species 


SUBGROUP I 


Hylephila herrerai MacNeill, new species 
(Figs. 4a, 12, 26, 32, 33, 58) 


Description. Male. Head. Dorsally scaled black with long, 
golden vestiture. Palpi and antennae missing. Body, collar, and pata- 
gia as head, dorsally with orange vestiture. Hind leg with two pairs 
of spurs. Wings. Stubby, rounded. Forewing length 9 mm (n = 1). 
Above stigma absent. Fulvous orange in discal cell (divided by lon- 
gitudinal fuscous streak), narrowly on costa through area of apical 
spots, broadly and slightly paler postmedially in space M1—M2, nar- 
rowly in space M2—M3, then broadly again in spaces M3—Cul and 
Cul-Cu2, very narrowly in distal half of space Cu2—2A, and a streak 
in anal cell; these spots outwardly prolonged along veins. Fringes 
basally fuscous, outer half whitish with very pale fulvous tint. Below 
fulvous sullied, much more extensive except fuscous in lower half of 
space Cu2—2A and all of anal cell. Hindwing above brown with ab- 
breviated fulvous postmedian macular band from veins Rs to Cu2 
with spot in space MI-M3 more than thrice width of other spots; an 
extremely narrow, almost obsolete ray-like streak through lower half 
of discal cell nearly to postmedian macular band in space M1—M3; 
fuscous border nearly cut to termen by narrow fulvous along veins. 
Fringes as on forewing but slightly darker. Below broadly orange- 
fulvous with paler whitish veins scarcely contrasting with fulvous 
spaces, and reduced black spots basally in lower costal cell, elongate 
in mid-space Sc—Rs, basal lower half in discal cell, and round-edged 
postmedial and border spots in spaces Rs-M1, M3-Cul, and 
Cul—Cn2; vannal fold fuscous and expanding into basal half of space 
Cu2-2A; veins 2A and 3A scarcely pale. Fringes whitish with fulvous 
tint. Genitalia. Eighth tergite (Fig. 4a) with lateral margin only 
slightly concave before caudal margin; terminal bristle-sockets 
scarcely enlarged caudad. Valva (Fig. 12) in lateral view with length 
of basal margin less than or subequal to one and one-half length of 
valva. Penis proximally not strongly curved dorsad, its length ex- 
ceeding length of genitalic capsule but slightly less than twice length 
of valva; titillators asymmetric, the left large and broadly thorn-like 


(7 (aq 
CoG 


Fic. 4. Eighth tergites in males of five species of Hylephila in 
left lateral view. Scale = 1 mm. a, H. herrerai, new species holo- 
type, CHILE, Parinacota, “Arica” Cotacotani, 4500 m, II-28-48 
(genitalia dissection # ¢ 6117-CDM), in CAS. b, H. pseudoher- 
rerai, new species holotype, PERU, Ayacucho, Reserva Nac. de 
Pampas Galeras, 4000 m, II-17-78, J. L.Venero (genitalic dissec- 
tion # 6 3820-JH), in MUSM. e, H. pseudoherrerai, new species 
paratype ¢ same data as holotype except date is I-27-78, (genitalic 
dissection # 6 6311-CDM), in MUSM. d, H. pallisteri, new species 
holotype, same specimen as in Fig. la. e, H. pallisteri, same spec- 
imen as in Fig. lb. f, H. tentativa, new species holotype, same 
specimen as in Fig. 1d. g, H. shapiroi, new species holotype, same 
specimen as in Fig, 2a. 


and sclerotically strapped to penis, the right narrowly thorn-like and 
not sclerotically strapped to penis; comuti asymmetric, the proximal 
large and bidentate, the distal evidently fragmented into two sube- 
qual, slender, unidentate thorns. Juxta with ventral caudal clefts about 
one-half length of juxta; separated median floor not nearly reaching 
caudal margin of juxta. Uncus in dorsal view somewhat rounded 
cephalad, gradually emarginate to pectines caudad; caudal cleft 
scarcely exceeding pectines cephalad; pectines minute, not anteri- 
orly rounded, each half broader than long, clearly arched in lateral 
view, the tines in dorsal view obscure, many, and scarcely separated. 
Gnathos in lateral view dorsally distinctly sclerotized, not exceeding 
pectines caudad, and greatly divergent ventrally from uncus. 
Female. Head and body as male. Antennal club large, about 
three-fourths length of shaft, club posteriorly black with scattered 
golden scales, anteriorly and below buffy, golden above; nudum 
brown, less than one-fourth length of club, shaft buff. Palpi with sec- 
ond segment shaggy with mixed orange and black, long, hair-like 
scales exceeding length of third segment which is scaled black and 
orange. Eyelash greater than one-half eye diameter, orange, and 
black scaled. Collar and patagia dorsally orange scaled with emer- 
gent, scattered, long, hair-like, black scales. Wings. Somewhat more 
produced and apically pointed than male (Fig. 33). Forewing length 
10 mm (n = 1). Above as male but fuscous markings more extensive 


VOLUME 56, NUMBER 2 


r 


e 


Fic.5. Eighth tergites in males of three species of Hylephila in 
left lateral view. Scale = 1 mm. a, H. galera ?, same specimen as in 
Fig. 2c. b, H. galera?, same specimen as in Fig. 2d. e, H. boulleti, 
CHILE, Antofagasta, Salar Aguas Calientes, 4000 m, II-27-60, L. 
Pefia (genitalic dissection # 6 6125-CDM), in AME. d, H. boulleti, 
ARGENTINA, Catamarca, Quebrada de los Cazadores, grupo aus- 
tral Nevadas del Aconquija, 5400-4800 m, XI-23-48, M. Lamb (gen- 
italic dissection # 5 6213-CDM), in CAS. e, H. peruana, same spec- 
imen as in Fig. 3d. 


in costal cell; below as male but fuscous marginal wedge-shaped 
spots more prominent from space M1—M2 to space R4—R5 where 
otherwise inconspicuous pale veins are conspicuous to termen. 
Hindwing above as male, fulvous increased discally in lower distal 
half of discal cell continuing ray-like through spaces M1—M3 and in 
lower half of spaces Cu2-2A and 2A— 3A; below as male but with 
broadly scaled white veins more prominent, fuscous spots reduced. 
Fringes whitish with fulvous tint vannally. Genitalia. (Fig. 26). 
Highth sternite not sclerotized. Apophysis anterioris in lateral view 
scarcely produced cephalad of junction with lamella postvaginalis. 
Lamella postvaginalis in ventral view broadly medially united, each 
half produced ventro-cephalad forming an anterior bulge not 
nipple-like, medially produced ventrad into a narrow, double-folded, 
U-shaped flange just caudad of the mostly membranous antrum. 
Antrum caudodorsally weakly and irregularly sclerotized, anterodor- 
sally weakly sclerotized immediately caudad of ductal constriction. 
Ductus bursae weakly sclerotized laterally, ventrally mostly mem- 
branous, left lateral pocket not produced. 

Types. Holotype ¢, CHILE, “Arica,” Parinacota, Cotacotani, 
4500 m, [1-28-48 (genitalic dissection # ¢ 6117-CDM), in CAS. 
Paratype. 1 2, same data as holotype but additional determination la- 
bel by K. J. Hayward as Hylephila phylaeus basistrigata Eaton. (gen- 
italic dissection # 2 6118-CDM), in CAS. 

Etymology. With considerable pleasure I name this species for 
my good friend, the late Prof. José Valentin Herrera G., to whom 


69 


this paper is dedicated. Pepe Herrera devoted well over four 
decades to the collection and study of Chilean Lepidoptera. He was 
actively involved with this study of Hylephila and co-authored our 
first paper on this genus. 

Diagnosis and discussion. This species is distin- 
guished by its small size, lack of a stigma in the male, 
and the broadly darkened vannal area above. It differs 
from males of H. pseudoherrerai in its smaller size, 
darker wing markings, and in genitalic details of the 
uncus, pectines, and juxta. The female genitalia most 
resemble those of H. tentativa in the lack of a sclero- 
tized eighth sternite, but the very short apophyses an- 
teriores, and weakly sclerotized, short ductus bursae 
differ from these structures in H. tentativa. The wing 
borders are not as broadly cut by fulvous along the 
veins as are those of H. tentativa. The species flies in 
February. 


Hylephila pseudoherrerai MacNeill, 
new species 
(Figs. 4b, 4c, 13, 14, 34, 35, 58) 


Description. Male. Head. Dorsally dark, black and golden scal- 
ing mixed equally, except for pale golden patch over eye behind 
black eyelash. Antennae with club black posteriorly, anteriorly buff 
merging to golden above, club about one-half length of shaft, 
nudum dark brown, about one-fourth length of club. Palpi golden, 
second segment somewhat shaggy, long, Bee hair-like scales re- 
stricted to anterolateral angles, third segment dorsally black, 
scarcely emerging from vestiture of second segment. Body. Dorsally 
black with long, golden, hair-like overscaling, except for pale g golden 
patagia and pale margins of otherwise black tegulae. Hind tibia with 
two pairs of spurs. Wings. Narrow and rounded to somewhat 
rounded at tornus. Forewing length, holotype 11 mm, one paratype 
12 mm. Above stigma absent. Fulvous rich to cold orange costad of 
discal cell merging with apical spots (but sullied basad), in discal cell 
(where divided by broad, blackish fuscous streak from base to end 
cell where connected broadly to elongate black spot between end 
cell and subterminal spots), and below cell in base of space 
Cul-Cuz2, in subterminal spots and in postmedian band from spaces 
M3—Cul to Cu2—2A, where sullied, and in most of cell where also 
sullied. Fuscous border broad, only slightly cut by fulvous veins. 
Fringes brownish, at tornus sullied fulvous. Below costad, apically 
on veins, and distally on fringes, whitish. Hindwings above with ful- 
yous somewhat restricted to a ray in distal half of cell through spaces 
M1-M3 to broad, fuscous border, in cell Cu2—2A from near base 
nearly or quite to margin, and narrowly as part of the postmedian 
macular band in spaces Cul—Cu2 and M3-Cul, and scarcely evi- 
dent in space Rs—M1; fuscous border broad, scarcely cut by fulvous 
veins. Fringes pale fulvous to vannally fulvous and apically fuscous. 
Below, extensively pale fulvous with contrasting white veins Sc to 
Cu2; black spots basally in costal cell, discal cell, and ray-like from 
base to margin in space 2A—3A and anal cell, postmedially near base 
of spaces Rs—M1, M3-Cul, and Cul—Cu2, indistinctly and ray-like 
in Sc+R1-Rs, marginally in spaces Rs—M1, M3-Cul, and Cul—Cu2. 
Fringes slightly paler than above with a slight violet tint. Genitalia. 
Highth tergite (Figs. 4b, 4c) lateral margins slightly emarginate be- 
fore caudal margin; terminal bristle-sockets slightly oalkveaa before 
caudal margin. Valva (Figs. 13, 14) narrow, in lateral view with 
length of basal margin more than (or slightly less than) one and one- 


half depth of valva. Penis strongly curved dorsad anteriorly (or 
straight), its length exceeding length of capsule and subequal to 
twice length of valva; titillators sclerotically strapped to penis, 
slightly asymmetric, the left more robust than the right, thorn-like 
with its point (more or less) angled ninety degrees from its base, cor- 
nuti bidentate, slightly asymmetric. Juxta with ventrocaudal clefts 
about one-half length of juxta, the separated median floor nearly 
reaching caudal margin of juxta. Uncus in dorsal view greatly ex- 
panded and rounded cephalad, then abruptly emarginate caudad, or 
smoothly tapered, to pectines; caudal cleft distinctly exceeding 
pectines cephalad; pectines small, each half broader than long, not 
anteriorly rounded, the tines conspicuous, few; in lateral view some- 
what arched. Gnathos in lateral view dorsally distinctly sclerotized, 
not to slightly exceeding pectines caudad, and greatly to somewhat 
divergent ventrad from uncus. 

Female. Unknown. 

Type. Holotype ¢, PERU, Ayacucho, Reserva Nac[iondl] de 
Pampas Galeras, 4000 m, 17-II-78, J. L. Venero (genitalic dissection 
# 3 3820-JH), in the Museo Nacional de Historia Natural, Pert 
(MUSM). Paratypes. 1 d, same locality and collector as holotype, but 
27-I-78, (genitalic dissection # 6 6311-CDM); 1 4, same locality but 
date 21-IV-74 and G. Lamas, collector (genitalic dissection # 5 3736- 
JH). Paratypes will be placed in MUSM and CAS. 

Etymology. This species, while small and without a male stigma, 
is not the same species as H. herrerai. 

Diagnosis and discussion. The holotype has nar- 
rower wings and is more orange than one paratype. 
The latter is larger, paler, and looks very much like a 
female of H. boulleti, which I thought it was until I dis- 
sected it. The genitalia are slightly different in the type 
and one paratype; those of the paratype (# ¢ 6311- 
CDM) somewhat resemble those of H. herrerai, but 
the locality data and the contrasting white veins on the 
hindwing venter suggest that all three specimens are 
samples of the same population. Hylephila herrerai is 
smaller with broader fuscous markings above and lacks 
contrasting white veins below, and the genitalia of H. 
herrerai differ in the narrower uncus, smaller pectines 
with inconspicuous tines, and a shorter juxtal median 
floor. The species has been collected in January, Feb- 


ruary, and April. 


Hylephila pallisteri MacNeill, new species 
(Figs. la, 1b, 4d, 4e, 15, 16, 36, 37, 58) 


Description. Male. Head. Dorsally scaled orange fulvous; eye- 
lash black with scattered orange hairs. Antennae with club anteriorly 
and above orange, merging to white below, posteriorly black; nudum 
orange shading to brown on apiculus; shaft length more than twice 
length of club. Palpi with third segment protruding above vestiture 
of second segment; second segment vestiture black above and on 
distal third below, lateral scales orange blending to white, hair-like 
scales anteriorly orange, but at anterolateral angles black. Body, 
patagia, and collar as head, dorsally with orange vestiture. Hind legs 
with two pairs of spurs. Wings. Broad. Forewing length 11 mm (sec- 
ond specimen 14 mm) (n = 2). Above stigma present, slender; mi- 
croandroconial mass black, inconspicuous amid other black stigmal 
elements; post-stigmal patch conspicuous, broad. Orange fulvous 
discally greatly reduced by broad fuscous border scalloped along 
veins, but broadly orange in costal and subcostal spaces from base 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


nearly to subapical fulvous spots, and through entire discal cell ex- 
cept where divided longitudinally by slender fuscous streak; narrow 
subterminal spots distinctly offset from only slightly broader post- 
median macular band. Fringes broadly fuscous, terminally pale ful- 
vous. Below, stigma pocket upper element extends well beyond ori- 
gin of vein Cul and diverges from discal cell about equidistant from 
origins of veins Cul and Cu2, lower elements large (Figs. la, 1b). 
Orange fulvous as above but broader discally and more restricted 
basad by fuscous in spaces Sc-R, basal half of Cu2—2A, discal cell, 
and entire anal cell; costa white and veins R3 to M2 distally white, 
black bar end discal cell and fuscous spots postmedially in spaces 
M1—M2, M2-M3, and M3-Cul more or less heavily overscaled with 
fulvous, and distal to stigma pocket in spaces Cul—Cu2 and Cu2—2a. 
Hindwing above fuscous, fulvous restricted to streak through lower 
half of discal cell and a very narrow postmedian macular band 
Cu2-2A to crescent in Rs—M1, and weakly along vein 2A to termen, 
fringes as forewing but more orange. Below fulvous with veins (ex- 
cept M2) from Cul to Sc strongly contrasting white; rounded black 
spots basally in costal and discal cells, postmedially and marginally in 
spaces Sc—Rs, Rs—M1, M3-Cul, Cul—Cu2, and discally in upper 
half of discal cell; space Cu2—2A fulvous from base to termen, vein 
3A distal two-thirds fulvous; fringes more orange than above. Geni- 
talia. Eighth tergite with lateral margin broadly emarginate before 
caudal margin (Figs. 4d, 4e), ), terminal bristle-sockets enlarged just 
before caudal margin. Valva (Figs. 15, 16) in lateral view narrow, 
basal margin not strongly convex, its length subequal to one and 
one-half times depth of valva, dorsal margin not greatly concave to 
caudal beak; in caudal view lower half of horizontal cleft narrow, 
scarcely one-half width of knobbed upper half. Penis proximally 
slightly curved dorsad, its length exceeding length of genitalic cap- 
sule and about twice length of valva; titillators asymmetric, the left 
large, broadly thorn-like, and sclerotically strapped to penis, the 
right much reduced, narrowly thorn-like, and obscurly strapped to 
penis; cornuti bidentate, asymmetric, the proximal basally much 
elongate, the distal not elongate. Juxta with ventrocaudal clefts 
short, about one-fourth length of juxta; separated median floor 
reaching caudal margin of juxta. Uncus in dorsal view distinctly 
round cephalad, abruptly tapered to caudal pectines; the caudal cleft 
not exceeding pectines cephalad; pectines small, each half longer 
than broad, anteriorly and posteriorly rounded, the tines few, con- 
spicuous, in lateral view scarcely arched. Gnathos in lateral view dor- 
sally clearly sclerotized, not exceeding pectines caudad, and greatly 
divergent ventrad from uncus. 


Female. Unknown. 

Type. Holotype ¢, PERU, Cuzco, Ollantaitambo, II-24-47, 9200 
ft., C. J. Pallister (genitalic dissection # 4 3811-JH), in AMNH. 

Etymology. This species is named for the collector of the holo- 
type, C. J. Pallister. 

Diagnosis and discussion. Two additional speci- 
mens have been \seen. One, a male from Yungas (or 
LaPaz) Bolivia, 1000 m, ex H. Rolle, 1902, R. 
Oberthur Coll’n. (genitalic dissection # ¢ 3815-JH) was 
prematurely returned to the BMNH identified as H. 
boulleti, but was noted as having identical genitalia to 
specimen # ¢ 3811-JH (the holotype of H. pallisteri). I 
have not examined this specimen in detail. 

A more recent specimen (Figs. 1b, 4e, 16, 37, 58) 
from PERU, Cuzco, Abra Acjanacu, 3600 m. 17-V-84, 
d. Lamas (genitalic dissection # 6 6227-CDM), in 
MUSM, seems to be this species; but it differs in sev- 


VOLUME 56, NUMBER 2 


eral particulars from the holotype: It is much larger 
with a forewing length of 14 mm. The antennal club is 
longer so the shaft is about twice the length of the 
club. The third palpal segment is ventrally entirely ful- 
vous, not black on the distal third. On the wings above 
the post-stigmal patch is not as broad as in the holo- 
type, and below the stigmal pockets are more robust. 
The markings are quite similar; but on the hindwing 
venter there is an extra elongate postmedian black spot 
in the upper fourth of space M1—M3, and vein M2 is 
apparent and whitened. The genitalia also differ in 
that the valva is a bit narrower with the length of the 
basal margin more than one and one-half times the 
depth, and the lower one-half of the horizontal beak 
cleft in caudal view is nearly as wide as the upper half. 
The penis is not as strongly bent dorsad proximally, 
and the discal cornutus is unidentate. The juxta has the 
ventrocaudal clefts long, about half the length of the 
juxta. The uncus in dorsal view has the lateral margin 
more abruptly emarginate caudad before the pectines 
and the caudal cleft distinctly exceeding the pectines 
cephalad. The pectines are small with each half 
scarcely longer than broad, and anteriorly as well as 
posteriorly scarcely rounded, and the tines are numer- 
ous and inconspicuous. The gnathos in lateral view is 
only somewhat divergent ventrally from the uncus. 
These characters suggest that perhaps this specimen 
represents a closely related species. 

The two specimens are distinctive in the orange ful- 
vous above with broad borders, the very black stigmal 
elements including the microandroconial mass, and 
the dark fuscous, orange fulvous, and contrasting 
white veins below. The specimens were collected in 
March and May. 


Hylephila blancasi MacNeill, new species 
(Figs. lc, 6, 9, 17, 38, 39, 58) 


Description. Male. Head. Dorsally pale fulvous. Antennal club 
anteriorly and above orange fading to pale buff below, posteriorly 
black; nudum pale brown; shaft slightly longer than twice length of 
club. Palpi shaggy, pale fulvous, second segment front has hair-like 
scales pale, anterolateral angles black, laterally scales pale, not hair- 
like, third segment dorsally black, scarcely emerging from vestiture 
of second segment. Body dorsally black with long, pale, golden, 
hair-like overscaling, except for golden patagia and margins of oth- 
erwise black tegulae. Hind tibiae with two pairs of spurs. Wings. 
Forewing produced, length 12 mm (range 11-13 mm) (n = 5). 
Above stigma present; microandroconial mass dark gray, inconspic- 
uous; post-stigmal patch narrow. Fulvous extensive from costa 
through discal cell, basally sullied, a short black streak dividing dis- 
tal half and an oblique black line at end of discal cell; fuscous border 
divided to termen by fulvous along veins; inner edge of subterminal 
fulvous spots usually in line with inner edge of postmedian macular 
band. Below, stigma pocket upper element extends just beyond ori- 
gin of vein Cul, diverges from discal cell cubitus vein much nearer 
to origin of vein Cul than Cu2, and reaches vein Cu2, lower ele- 


blantasi 


Fic. 6. SEM image of male eighth tergite (dorsal aspect) of H. 
blancasi, new species paratype (descaled to show caudal array of 
bristle-sockets at 70x, with inset enlarged below). [PERU, Lima] 
cerca (sic) Canta, 4700 m, XII-13-47, Coll. Blancas (SEM #9, 23- 
CDM), in CAS. 


ments very large (Fig. lc); costa whitened (gray where worn) and 
veins Rs to M3 buff distally (clearly white in two paratypes); exten- 
sive fulvous pale, obscuring subapical fulvous spots; discal cell, costal 
cell, and space Sc-R all black at base, as is basal half cf space 
Cu2-2A and all of anal cell; marginal fuscous spots isolated by pale 
fulvous along veins to termen, that in space MI—M2 nearly obsolete; 
diagonal black bar at end of discal cell, fuscous postmedian spots in 
spaces Cul—Cu2, M1—M2, and M2-M3, the latter two heavily over- 
scaled fulvous. Hindwings above with costal cell and space Sc—Rs 
fuscous, fuscous border nearly or quite divided to termen by fulvous 
extensions along veins, fulvous of space MI—M3 a ray-like extension 
well into discal cell, fuscous basal half of postmedian spaces 
Cu2-2A, Cul—Cu2, and discal cell all heavily overscaled with ful- 
vous; postmedian fulvous spot in space Rs—M1 an angular crescent, 
the upper half twice as wide as lower half. Below pale fulvous re- 
stricted by white scaling very broadly along side of veins Rs to Cu2, 
but scales of veins M3, Cul, and Cu2 light brownish, contrasting 
with whitish to pale fulvous intervein spaces (one paratype has thin 
white streak mid fulvous of space Cu2—2A), space 2A—3A and anal 
cell wholly fuscous, vein 1A a thin fulvous streak, basal and post- 
medial black spots edged or partly overscaled by cupreous scales or 
hairs. Fringes dusky, vannally pale orange. One paratype (# ¢ 3819- 
JH) has fulvous extensive and dark markings much reduced like H. 
galera and forewing subterminal fulvous spots in spaces M1—M2 and 
M2-M3 slightly offset from postmedian band spots in spaces 
M3-Cu2 to Cu2-2A owing to fulvous expanded basad in M3-Cul. 
Another paratype (SEM # ¢ 23-CDM) has more extensive dark 


bouletti 


Fic. 7. SEM image of male eighth tergite (dorsal aspect) of H. 
boulleti (descaled to show caudal array of bristle-sockets at 70x with 
inset enlarged below). CHILE, Parinacota, nr. base Vol. Guallatire, 
4500 m, XI-21-94, A. M. Shapiro (SEM #7-CDM), in CAS. 


markings above so the fulvous subterminal spots are not only offset 
but completely separated from the fulvous postmedian band. Geni- 
talia. Eighth tergite with lateral margin distinctly emarginate before 
caudal margin, terminal bristle-sockets conspicuously enlarged cau- 
dad, more or less rounded in cross section (Fig. 6). Valva (Fig. 17) in 
lateral view broad, with ventral margin strongly convex but distad 
abruptly concave to a prominent beak; dorsal margin basally 
abruptly concave to the caudal beak, length basal margin slightly 
greater than one and one-half times depth of valva. Penis proximally 
scarcely (or not) curved dorsad, its length exceeding length of geni- 
talic capsule and greater than twice length of valva; titillators nearly 
symmetric, broadly based, thorn-like, and sclerotically strapped to 
penis, the right slightly reduced; cornuti asymmetric, bidentate, one 
much more broadly elongate. Juxta with ventral caudal clefts long, 
about one-half length of juxta, separated median floor scarcely 
reaching caudal margin of juxta. Uncus in dorsal view with distinctly 
ourndled cephalic and lateral mar gins, then emarginate and abruptly 
tapered, without serrations, to caudal pectines, the caudal cleft not 
(or slightly) exceeding pectines cephalad; pectines minute (Fig. 9), 
each half about as long as broad, anteriorly and posteriorly not, to 
somewhat, rounded, the tines many, inconspicuous, somewhat 
arched in lateral view. Gnathos massive in lateral view, dorsally dis- 
tinctly sclerotized, not exceeding pectines caudad, and greatly diver- 
gent ventrally from uncus. 

Female. Unknown. 

Type. Holotype 3, PERU, Santia, donated 1902, P. forte [sic] 
(genitalia dissection # d 3873-JH), in AMNH. Paratypes. 1 }, 
[PERU, Lima] cerca Canta, 4700 m, 13-XII-47, Coll. Blancas (SEM 
#3 9, 23-CDM); 1 6, [PERU, Junin] entre La Oroya ane 10-I- 
49, Coll. Blancas (genitalic dissection # ¢ 3819- -JH); 1 d, [PEri] 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fic. 8. SEM image of male eighth tergite (dorsal aspect) of H. 
peruana (descaled to show caudal array of bristle-sockets at 70x, 
with inset enlarged below). PERU, Junin, Pachachaca, 4000 m, V- 
20-79, G. Lamas (SEM #8-CDM), in MUSM. 


#22694, E. E. Olcott (genitalic dissection # 5 3874-JH); 1 4, [PERU] 
illegible locality, Huanuco, Holland Collection (genitalic dissection 
# 3 6280-CDM). Paratypes will be placed in the following collec- 
tions: AMNH, CAS, CMNH. 

Etymology. This species is named for the collector of the first 
two specimens I saw of this species. The holotype was selected be- 
cause it is in better condition. 


Diagnosis and discussion. Hylephila blancasi has 
superficial markings similar to those of H. galera. One 
paratype (genitalic dissection # 3 3819-]H) is very like 
the latter with pale fulvous extensive on the dorsal 
forewing, the dark markings much reduced, and the 
forerting subterminal fulvous spots in spaces M1—M2 
and M2_M3 slightly offset from the postmedian band 
of fulvous spots from M3-Cul to Cu2—2A, owing to 
the fulvous expanded basad in space M3—Cul. Hyle- 
phila galera, however, has much broader wings and 
genitalically belongs with subgroup II. Another 
paratype of H. blancasi (SEM # 6 9, 23-CDM) has the 
dorsal forewing subterminal spots reduced and com- 
pletely separated from a narrowed postmedian band 
by a combination of a distally expanded subapical fus- 
cous spot and a greatly broadened marginal border. 
Hylephila blancasi flies in December and January. 

The following species, H. tentativa, is remarkably 
similar to H. blancasi; but the fulvous of the wing 


VOLUME 56, NUMBER 2 


Fic. 9. SEM image of caudal tip of uncus (dorsal aspect) of 
male H. blancasi showing pectines at 400x. Same specimen as Fig. 6 
(SEM #9, 23-CDM), in CAS. 


above is much more orange on H. tentativa, the male 
stigma is less well developed, and the genitalia are 
somewhat different. The valva of H. blancasi is much 
broader, the titillators are more robust and the cornuti 
distinctly bidentate, the juxta is less robust, the uncus 
in dorsal view is anteriorly more rounded and the 
pectines slightly larger with numerous, inconspicuous 
tines, and the gnathos is caudally more or less incon- 
spicuous from above than are these structures in H. 
tentativa. 


Hylephila tentativa MacNeill, new species 
(Figs. 1d, 4f, 18, 27, 40, 41, 58) 


Description. Male. Head. Dorsally black with mixed golden 
and black hair-like vestiture. Antennae dorsally black on club, black 
mixed with white scales on shaft, anteriorly golden on club and shaft 
fading to buff basally; club less than one-half length of shaft, nudum 
pale brown; shaft length slightly greater than dorsal width of head. 
Palpi shaggy; third segment black, slightly protruding anteriorly 
from vestiture of second segment in dorsal view, in lateral and ven- 
tral views scarcely protruding from rich, golden, fulvous, hair-like 
vestiture of front of second segment. Eyelash length about one-half 
eye diameter. Body dorsally black with sparse golden, hair-like 
scales. Tegulae black with broad, pale edges. Legs with hind tibiae 
bearing two pairs of spurs. Wings. Somewhat narrow, slightly pro- 
duced. Forewing length 12 mm (n = 1). Above stigma present, evi- 
dent but not conspicuous; microandroconial mass very narrow, 
scarcely distinguishable between very abbreviated brush patches, 


boulleti 


Fic. 10. SEM image of caudal tip of uncus (dorsal aspect) of 
male H. boulleti showing pectines at 400x. CHILE, Parinacota, Re- 
serv. Nac. Salar de Surire, no. shore, 4250 m, XI-22-94, A. M. 
Shapiro (SEM #22-CDM), in CAS. 


Peruana 


Fic. 11. SEM image of caudal tip of uncus (dorsal aspect) of 
male H. perwana showing pectines at 400x. PERU, Podecayo, 
Feb.—May 1925 (SEM #24-CDM), in LACM. 


74 


Nu, 
Ny 


Fic. 12. Male genitalia of H. herrerai, new species holotype. 
Uncus (lateral and dorsal aspects, dorsal of pectines enlarged), val- 
vae (left outer and caudal aspects, right inner aspect), juxta (left lat- 
eral and ventral aspects), penis with vesica everted (left lateral and 
distal dorsal aspects). Data as Fig. 4a. Scale = 1 mm. 


post-stigmal patch narrow but most conspicuous stigmal element. 
Fulvous warm, ruddy, almost orange, abruptly contrasting with fus- 
cous markings; border cut to termen by fulvous along veins. Inner 
edge of fulvous subterminal spots not in line with inner edge of post- 
median macular band. Fringes basally fuscous, terminally buffy or- 
ange. Below, rich fulvous more extensive, fuscous markings reduced, 
veins R5, M1, and M2 whitened as is the costa from base to apex. 
Stigma pocket with upper element very narrow, its width scarcely 
twice diameter of adjacent cubitus vein, lower elements reduced, 
widely separated. Hindwing above with rich fulvous postmedian 
macular band from spaces R5-M1 to Cul—Cu2, and fulvous rays 
from base discal cell to margin in space MI-M3 and near base to 
margin in Cu2—2A; fuscous border cut to termen by fulvous along 
veins. Below fulvous somewhat paler but not nearly whitish, fuscous 
spots broadly separated by fulvous flanking the whitened veins from 
Rs to Cu2, space Cu2-2A from base to termen all fulvous, a poorly 
defined, narrow, pale ray from near base to termen along vein 3A in 
otherwise fuscous vannal area. Genitalia. Eighth tergite (Fig. 4f) 
laterally conspicuously emarginate immediately cephalad of caudal 
margin; terminal bristle-sockets conspicuously enlarged caudad. 
Valva (Fig. 18) in lateral view narrow, length basal margin distinctly 
more than one and one-half times valva depth, dorsal margin some- 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 13. Male genitalia of H. pseudoherrerai, new species holo- 
type. Uncus (lateral and dorsal aspects, dorsal of pectines enlarged), 
valvae (left outer and caudal aspects, right inner aspect), juxta (left 
lateral and ventral aspects), penis with vesica everted (left lateral and 
distal dorsal aspects). Data as Fig. 4b. Scale = 1 mm. 


what concave, caudal beak not prominent. Penis about twice length 
of valva; titillators sclerotically strapped to penis, slender, thorn-like, 
asymmetric, the left basally much more massive than the right; cor- 
nuti asymmetric, one bidentate and basally massive, the other 
smaller and minutely tridentate. Juxta with ventral caudal clefts 
about one-half length of juxta and separated mid-ventral floor nearly 
reaching caudal margin of juxta. Uncus in dorsal view more or less 
triangular, cephalic margin somewhat convex, and lateral margins 
from cephalic lateral angles only slightly sinuate and tapered to 
pectines, caudal cleft distinctly exceeding the pectines cephalad; 
pectines minute, not anteriorly rounded, each half longer than broad, 
the tines few, conspicuous, in lateral view scarcely arched. Gnathos 
massive, sclerotized, projecting caudad well beyond pectines in dorsal 
view, divergent from uncus in lateral view. 

Female. Head and body as male but fulvous paler, cheeks and 
eye ring white. Antennal club slightly less than one-half length of 
shaft, club black, posteriorly buffy, anteriorly and below, not check- 
ered on shaft; nudum brown, slightly less than one-half length of 
club. Palpi second segment shaggy, pale to buffy fulvous, black hair- 
like scales restricted to anterolateral angle. Eyelash about one-half 
eye diameter. Patagia dorsally fulvous. Wings. Somewhat produced 
and apically pointed. Forewing length 12.5 mm (n = 1). Above as 
male, fulvous paler, less ruddy, fuscous of discal “false stigma” more 


VOLUME 56, NUMBER 2 


14 


Fic. 14. Male genitalia of H. pseudoherrerai, new species, 
paratype. Uncus (lateral and dorsal aspects, dorsal of pectines en- 
larged), valvae (left outer and caudal aspects, right inner aspect), 
juxta (left lateral and ventral aspects), penis with vesica everted, (left 
lateral and distal dorsal aspects). Same specimen as in Fig. 4c. Scale 
= 1mm. 


extensive than male, border sharply cut to termen along veins from 
M1 to 2A, inner edge of subterminal fulvous spots not in line with 
inner edge of postmedian band; termen fuscous; fringes basally fus- 
cous, terminally paler except apically. Below pale fulvous as male but 
veins less whitened. Hindwing above as male but fulvous less ruddy. 
Below as male but fuscous markings slightly broader; termen fus- 
cous; fringes basally fuscous, terminally buffy. Genitalia. (Fig. 27). 
Highth sternite not sclerotized. Apophysis anterioris in lateral view 
produced cephalad of junction with lamella postvaginalis, its length 
subequal to sclerotized dorso-ventral width of papillae anales. 
Lamella postvaginalis in ventral view medially united, anteriorly pro- 
duced ventrad into a double-folded U-shaped flange, medially scle- 
rotically extended cephalad toward dorsal portion of antrum. 
Antrum dorsally and anterolaterally well sclerotized, ventrally dark- 
ened, plicate-membranous. Ductus bursae well sclerotized, left lat- 
eral pocket clearly produced. 

Types. Holotype 3 PERU, Ay[acucho], Apacheta de Tambo, 
4250 m, 12°59’S, 74°05’ W, 29-I-99, G. Lamas (genitalic dissection # 
3 6308-CDM), in MUSM. Paratype 1 ° PERU, Aylacucho], Rio 
Apacheta, 4200 m, 13°21’S, 74°39’W, 24-I-99, G. Lamas (genitalic 
dissection # 2 6309-CDM), in MUSM. 


=I 


Ol 


15 


Fic. 15. Male genitalia of H. pallisteri, new species holotype. 
Uncus (lateral and dorsal aspects, dorsal of pectines enlarged), val- 
vae (left outer and caudal aspects, right inner aspect), juxta (left lat- 
eral and ventral aspects), penis with vesica everted, (left lateral and 
distal dorsal aspects). Data as Fig. la. Scale = 1 mm. 


Etymology. The specific name reflects the uncertainty expressed 
in the discussion below. 


Diagnosis and discussion. As mentioned, this 
species is remarkably like H. blancasi; in fact when I 
first saw this pair of specimens I was convinced they 
were that species. I was surprised by the apparent dis- 
junction of this population from the rather more 
northern ones of the known H. blancasi. In comparing 
seemingly slight differences in color pattern, stigmal 
characters, and especially genitalia, I suspected that 
this was not H. blancasi; and since this pair seems to 
be a male and a female from the same population, I 
elected to emphasize this, but not without some hesi- 
tation—hence the name. The types were collected in 
January. 

The differences from H. blancasi in the male geni- 
talia are discussed under that species. The female gen- 
italia resemble those of H. herrerai in lacking a sclero- 
tized eighth sternite, but they differ from those of that 


76 


Fic. 16. Male genitalia of H. pallisteri? Uncus (lateral and dor- 
sal aspects, dorsal of pectines enlarged), valvae left outer and caudal 
aspects, right inner aspect ), juxta (left lateral and ventral aspects), 
penis with vesica everted (left lateral and distal dorsal aspects). Same 
specimen as in Fig. 1b. Scale = 1 mm. 


species in the length of the apophysis anterioris and in 
the clearly produced lateral pocket of the ductus bur- 
sae. There is scarcely any resemblance to that species 


superficially, 
SUBGROUP II 


Hylephila shapiroi MacNeill, new species 
(Figs. 2a, 4g, 19, 42, 58) 


Hylephila ? n. sp., cf. boulleti group, Shapiro, 1985. 
Stud. Neotrop. Fauna and Environ. 20: 9, 10, figs. 
5(3), 6(3). 

Description. Male. Head. Dorsally shaggy with golden hairs. 
Antennae anteriorly buff, posteriorly black; club about one-half 
length of shaft; nudum orange-brown; apiculus very short, appar- 
ently one abbreviated segment, its width greater than its length. 
Palpi, second segment shaggy with pale fulvous hairs, hairs black at 
anterolateral angles; third segment dorsally black-hirsute, ventrally 
fulvous, not exceeding hairs of second segment. Body. dorsally and 
ventrally black with golden hairs. Tegulae black with broad white 
edges. Legs with femora black, fringed with golden hairs, tibiae and 
tarsae white; hind tibia with single pair of spurs. Wings. Produced, 
narrow. Forewing length 8.5 mm (n = 1). Above stigma present but 
very obscure except under magnification, microandroconial mass 
gray, not nearly extending to origin of vein Cul, brush patches and 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


17 


Fic. 17. Male genitalia of H. blancasi, new species holotype. 
Uncus (lateral and dorsal aspects, dorsal of pectines enlarged), val- 
vae (left outer and caudal aspects, right inner aspect), juxta (left lat- 
eral and ventral aspects), penis with vesica everted, (left lateral and 
distal dorsal aspects). Data as Fig. lc. Scale = 1 mm. 


post-stigmal patches of erect scales absent. Costa white. Broadly fus- 
cous, with extremely pale, nearly hyaline, fulvous spots before end of 
discal cell, and subapical in spaces R4-R5 and R5—M1, in subtermi- 
nal spots, (which are nearly continuous with postmedian macular 
band, not offset), in postmedian band from M1I—M2 to Cul—Cu2; 
spots of band in Cu2-2A extremely pale fulvous, not nearly hyaline; 
basad fuscous overscaled fulvous either side of fuscous streak divid- 
ing discal cell and costad of discal cell to end cell. Fringes fuscous. 
Below stigma pocket robust but short, upper element not nearly ex- 
tending to origin of vein Cul, diverging from discal cell cubitus vein 
much closer to vein Cu2 than to vein Cul; lower elements tiny, up- 
permost broad (Fig. 2a). Spots as above but fuscous more exten- 
sively overscaled fulvous everywhere except in spaces Cul—Cu2, 
Cu2-2A, and anal cell; entire costa and veins R4 to M1 distally 
white. Hindwings above wholly fuscous except for extremely pale 
fulvous, nearly hyaline, postmedian macular band from space 
Rs—M1 to M3-Cul (in M1—M3 elongate and ray-like from end dis- 
cal cell) and pale fulvous in space Cul—Cu2. Fringes fuscous basally, 
white vannally and terminally. Below, black spots basally in costal 
and discal cells and space Cu2—2A; postmedially in spaces Sc—Rs, 
Rs-M1, Cul—Cu2, and end discal cell; border fuscous spots from 
distal end spaces Sc—Rs to Cul—Cu2; space 2A—3A entirely fuscous 
overscaled fulvous and anal cell black; veins from Sc to Cu2 white; 


VOLUME 56, NUMBER 2 


Fic. 18. Male genitalia of H. tentativa, new species holotype. 
Uncus (lateral and dorsal aspects, dorsal of pectines enlarged), val- 
vae (left outer and caudal aspects, right inner aspect), juxta (left lat- 
eral and ventral aspects), penis with vesica everted, (left lateral and 
distal dorsal aspects). Data as Fig. 1d. Scale = 1 mm. 


costa, distal half discal cell, and anterior half of distal two-thirds 
space Cu2-2A white; posterior half of distal two-thirds space 
Cu2-2A very pale fulvous, as are the distal portions of pale spots of 
spaces Rs—M1 to Cul—Cu2,- that in MI—M3 a streak centrally for 
distal two thirds, a buff spot discally in space Sc—Rs. Fringes both 
wings sullied white, white vannally. Genitalia. Eighth tergite (Fig. 
4g) with lateral margin not distinctly emarginate before caudal mar- 
gin; terminal bristle-sockets slightly enlarged before caudal margin. 
Valva (Fig. 19) in lateral view broad, with basal, dorsal, and ventral 
margins more or less convex, caudal beak not prominent; length 
basal margin much less than one and one-half depth of valva. Penis 
proximally reflexed slightly ventrad, its length greatly exceeding 
length genitalic capsule and more than twice that of valva, elongate 
ventrocaudally as a terminally rounded, spatulate “floor’; titillators 
nearly symmetric, huge, laterally compressed thorns, each termi- 
nally with slender hooks laterad, sclerotically strapped to penis; cor- 
nuti asymmetric, minutely bidentate. Juxta severely damaged, not 
definable. Uncus in dorsal view, anterior half rounded, posterior half 
parallel-sided, where scarcely minutely serrate to pectines; pectines 
large, each half medially longer than broad, anteriorly tapered to 
cleft, posteriorly truncate, the tines numerous, inconspicuous, in lat- 
eral view pectines dorsally arched. Gnathos in lateral view mostly 
membranous, narrowly sclerotized dorsad, not divergent nor ex- 
ceeding pectines caudad, but uncus caudad and pectines slightly 
arched dorsad from gnathos. 

Female. Unknown. 

Type. Holotype 3 PERU, Dept. Junin, vic. Abra Anticona, 4843 
m, X-19-83, A. M. Shapiro (genitalic dissection # 5 5005-JH), in 
CAS. 


19 


Fic. 19. Male genitalia of H. shapiroi, new species holotype. 
Uncus (lateral and dorsal aspects, dorsal of pectines enlarged), val- 
vae (left outer and caudal aspects, right inner aspect), penis with 
vesica everted, (left lateral and distal dorsal aspects). Data as Fig. 2a. 
Scale = 1 mm. 


Etymology. I am delighted to name this remarkable little species 
for Arthur M. Shapiro, the collector of the only known specimen. 
Art has provided enormous and continuous support throughout 
these studies with years of discussion and careful collecting in South 
America. 

Diagnosis and discussion. This small species dor- 
sally appears to be more black and white rather than 
black and fulvous, and it does not seem to have a 
stigma until it is examined under magnification. The 
single pair of spurs on the metathoracic tibiae is a 
unique characteristic of this species within the boulleti 
group of Hylephila. This specimen was figured by 
Shapiro (1985) in his paper summarizing behavioral 
and ecological observations of the high altitude 
pierines he studied. In that paper, he discussed the 
morphospecializations of the genus Phulia Herrich- 
Schiffer and suggested that the convergence of the 
high altitude hesperiid fauna to the “dashed Phulia 
pattern” of high Andean pierines was owing to adapta- 


20 


Fic. 20. Male genitalia of H. galera? Uncus (lateral and dorsal 
aspects, dorsal of pectines enlarged), valvae (left outer and caudal 
aspects, right inner aspect), juxta (left lateral and ventral aspects), 
penis with vesica everted, (left lateral and distal dorsal aspects). Data 
as Fig. 2c. Scale = 1 mm. 


tion for crypsis. The type was collected in October fly- 
ing with H. peruana. 


Hylephila galera Evans 
(Figs. 2b, 2c, 2d, 5a, 5b, 20, 21, 43, 44, 45, 58) 


Hylephila galera Evans, 1955. Cat. Amer. Hesp. part 
IV, p. 314, pl. 75. Bridges, 1983, Lepid.:Hesp. Notes 
on species group names, part 1:47. 

In the description below, brackets, indicate remarks 

not taken from the type but from specimens #6121 

and/or #6307. 


Description. Male. Head. Dorsally black with orange hairs; 
antennae with club about one-half length of shaft, anteriorly buff 
merging dorsally to fulvous and ventrally to white, posteriorly black 
with scattered fulvous to white scales, nudum orange-brown, apicu- 
lus with ultimate segment slightly longer than wide; palpi missing 
[#6121: palpi with third segment evident but scarcely exceeding 
long, hair-like vestiture of second segment]. Body. [#6121: dorsally 
black with long golden hairs, ventrally black with long, shaggy, ful- 
vous hairs. Legs black, overscaled buff, with long fringes of orange 
hairs, hind tibiae with two pairs of spurs]. Wings. Broad, rounded. 
Forewing length 13 mm [range 13-14 mm] (n = 3) Above, stigma 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 21. Male genitalia of H. galera? Uncus (lateral and dorsal 
aspects, dorsal of pectines enlarged), valvae (left outer and caudal 
aspects, right inner aspect), juxta (left lateral and ventral aspects), 
penis with vesica everted, (left lateral and distal dorsal aspects). Data 
as Fig. 2d. Scale = 1 mm. 


conspicuous but narrow, apical and lower brush patches evident, as 
is the rather narrow post-stigmal patch [#6121: microandroconial 
mass dark brown]. Fulvous extensive, conspicuously penetrating 
wide fuscous border along veins to termen, inner edge subterminal 
fulvous spots offset from inner edge postmedian macular band; 
fringes basally fuscous, terminally orange. Below, stigma pocket 
narrow, upper element extends proximad from just beyond origin of 
vein Cul, diverges from discal cell cubitus vein much closer to ori- 
gin of vein Cul than to Cu2, and does not nearly reach vein Cu2; 
lower elements well separated from veins Cu2, 2A, and from each 
other (Fig. 2b). [#6121: upper element stigma pocket extends just 
beyond origin vein Cul and diverges from discal cell cubitus vein 
about equidistant from origins veins Cul and Cu2, without quite 
reaching vein Cu2; lower elements small, well separated (Fig. 2c); 
#6307: upper element nearly reaching vein Cu2, lower elements 
large, not widely separated (Fig. 2d )]. Fulvous extensive, costa and 
veins not whitened [#6121, 6307: veins R4 to M1 and costa 
whitened, at least under magnification], but veins R5 and M1 much 
paler distad. Hindwings above fulvous as on forewings penetrating 
border to termen. Below rich fulvous very extensive, entering lower 
half of discal cell nearly or quite to base, none of the veins whitened 
[#6121, 6307: veins of discal cell and from Rs to Cu2 white under 
magnification]. Fringes both wings basally sullied orange [#6121: or 
fuscous], terminally orange [#6121: or white]. Genitalia. Badly 
damaged; uncus eaten, penis, vinculum partly eaten, saccus miss- 
ing, valvae damaged, titillators and cornuti intact but not everted. 
[#6121 6307: eighth tergite lateral margin broadly concave and 
abruptly emarginate before caudal margin (Figs. 5a, b). Valvae 
(Figs. 20, 21) in lateral view somewhat broad, ventral and basal 


VOLUME 56, NUMBER 2 


Fic. 22, Male genitalia of H. boulleti. Uncus (lateral and dorsal 
aspects, dorsal of pectines enlarged), valvae (left outer and caudal 
aspects, right inner aspect), juxta (left lateral and ventral aspects), 
penis with vesica everted (left lateral and distal dorsal aspects). Data 
as Fig. 5c. Scale =] mm. 


margins gently convex, dorsal margin and ventral margins caudally 
concave to caudal beak; length basal margin about one and one-half 
times depth of valva. Penis proximally curved dorsad, its length 
subequal to length of genitalic capsule and almost twice length of 
valva; titillators short, thorn-like, sclerotically strapped to penis; 
cornuti nearly symmetric, very narrowly bidentate. Juxta with ven- 
trocaudal clefts long, nearly one-half length of juxta. Uncus in dor- 
sal view roughly triangular, not distinctly rounded cephalad, later- 
ally abruptly angled posteriorly one-half distance to very gradually 
tapered, laterally serrate, posterior half to caudal pectines, the cau- 
dal cleft not or scarcely exceeding pectines cephalad; pectines 
large, each half slightly longer than broad, anteriorly rounded, pos- 
teriorly emarginate, the tines numerous, inconspicuous. Gnathos in 
lateral view scarcely sclerotized, not exceeding pectines caudad, 
and not divergent ventrally from uncus]. 

Female. Unknown. 

Type. Holotype 6 PERU, Dept. Junin, Galera Pass, 4800 m, II- 
?-00 (Simons), “snowy season,” in BMNH. 

Discussion. The two additional specimens I have 
seen I believe represent this species, but they differ 
somewhat from the type, most particularly in the ele- 
ments of the stigma pockets. Neither has the upper el- 
ement as reduced as does the type, and # d 6307 has 


the lower elements very much larger and proximate 


79 


23 


Fic. 23. Male genitalia of H. boulleti. Uncus (lateral and dorsal 
aspects, dorsal of pectines enlarged), valvae (left outer and caudal 
aspects, right inner aspect), juxta (left lateral and ventral aspects), 
penis with vesica everted (left lateral and distal dorsal aspects). Data 
as Fig. 5d. Scale = 1 mm. 


(Fig. 2d). These specimens will be placed in the fol- 
lowing collections: #6121, from Tarma, Junin, Pert, 
3000 m, in LACM, and #6307, from Yauli, Corpacan- 
cha, Junin, Pert, 4300 m, in MUSM. This species flies 
in January and February. 


Hylephila boulleti (Mabille) 
(Figs. 8a, 3b, 5c, 5d, 7, 10, 22, 23, 28, 29, 46, 47, 48, 
49, 50, 51, 59) 


Chaerephon boulleti Mabille, 1906. Bull. Soc. Ento- 
mol. France, no. 6:67—68. 

Hylephila boulleti, Ureta, 1956. Bol. Mus. Nat. Hist. 
Nat. 26: 176, pl. 2, fig. 5b—5d; MacNeill & Herrera, 
1999, J. Lepid. Soc. 52:291. 

Hylephila boulletti (sic) boulleti, Ureta, 1963. Bol. 
Mus. Nac. Hist. Nat. 28:78,79. 

Hylephila boulleti boulleti, Evans, 1955. Cat. Amer, 
Hesp. part IV, p. 314, pl. 755; Hayward, 1973. 
Opera Lilloana 23:93; Lewis, 1973, Butterflies of 
the World, p. 246, pl. 83, fig. 17; Bridges, 1983, 


80 


24 


Fic. 24. Male genitalia of H. rossi, new species holotype. Uncus 
(lateral and dorsal aspects, dorsal of pectines enlarged), valvae (left 
outer and caudal aspects, right inner aspect), juxta (left lateral and 
ventral aspects), penis with vesica everted (left lateral and distal dor- 
sal aspects). Data as Fig. 3c. Scale = 1 mm. 


Lepid.: Hesp. Notes on species group names, part 

1:18; Shapiro, 1993, J. Res. Lepid. 30:163, 164; Pefia 

& Ugarte, 1997, Las mariposas de Chile, the butter- 

flies of Chile, p. 128, figs. 

Description. Male. Head. Dorsally very pale fulvous, with 
long, black hair-like scales rising from paired, postmedial black scale 
patches which are separated by smaller, triangular, medial patch on 
the vertex; pale, fulvous hair-like scales elsewhere except for mixed 
black and fulvous eyelash scales. Antennae anteriorly pale buffy to 
white, posteriorly black; club about one-half length of shaft; nudum 
pale orange to orange brown on apiculus; apiculus about three short 
segments, its basal width about equal to its length. Palpi third seg- 
ment black, ventrally mostly pale fulvous to buff, clearly to scarcely 
emergent from shaggy vestiture of second segment, where antero- 
lateral angles have scattered black hairs. Body dorsally black with 
scattered, long, pale golden vestiture of hair-like scales concentrated 
medially, pectus and venter white to pale fulvous. Legs mostly white, 
fringed fulvous, hind tibiae with two pairs of spurs, the upper occa- 
sionally somewhat reduced. Wings. Somewhat pointed apically, not 
stubby. Forewing average length 10.7 mm [range 9-13 mm] (n = 40). 
Above, stigma slender, conspicuous, microandroconial mass incon- 
spicuous without magnification, gray or tan to (rarely) yellow, apical 
and lower brush patches present, poststigmal patch conspicuous but 
not broad. Fulvous pale tawny, very variable in extent (cf. Figs. 46 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 25. Male genitalia of H. peruana. Uncus (lateral and dorsal 
aspects, dorsal of pectines enlarged), valvae (left outer and caudal 
aspects, right inner aspect), juxta (left lateral and ventral aspects), 
penis with vesica everted, (left lateral and distal dorsal aspects). Data 
as Fig. 3d. Scale = 1 mm. 


and 48); fuscous border if broad and clearly cut by fulvous veins, the 
fulvous not reaching termen along veins. Below, stigma pockets slen- 
der (Figs. 3a, b), diverging from discal cell cubitus vein about 
equidistant from origins of veins Cul and Cu2 or usually slightly 
nearer to that of vein Cul (but see Fig. 3b). Fulvous more extensive, 
costa and most veins whitened distally to termen. Fringes pale, basal 
third fuscous, middle third light fulvous, terminal third white. Hind- 
wings above as forewings, variable, border broad or narrow, usually 
not cut to termen by fulvous along veins. Postmedian macular band 
conspicuous, usually extending ray-like into at least lower part of dis- 
cal cell from space M1—MB3 and from vein 2A to Rs. Vein 2A fulvous 
from base to termen defining vannal fuscous area. Below pale ful- 
vous to almost white. Costa and all veins whitened, often including 
vein 3A, vannal area fuscous. Genitalia. Eighth tergite (Figs. 5c, d) 
with lateral margin broadly concave or sinuate cephalad, not dis- 
tinctly emarginate just before caudal margin; terminal bristle-sockets 
rounded in cross section and enlarged before caudal margin (Fig. 7). 
Valva (Figs. 22, 23) in lateral view broad, ventral margin convex, 
basal margin straight, dorsal margin slightly concave to caudal beak 
where knobs dorsad of caudal horizontal cleft few; length basal mar- 
gin about equal to one and one-half depth of valva. Penis proximally 
slightly curved dorsad, its length slightly greater than length of gen- 
italic capsule and less than twice length of valva; titillators large, 
asymmetric, the left very broadly and the right more narrowly thom- 
like, both sclerotically strapped to penis; cornuti asymmetric, one 
long, narrow, and basally very elongate, the other short, thorn-like, 
both minutely bidentate. Juxta broad, not elongate, with ventrocau- 
dal clefts long, about one-half length of juxta, separated median floor 
nearly or quite reaching caudal margin of juxta. Uncus in dorsal view 


VOLUME 56, NUMBER 2 


Fic. 26. Female genitalia of H. herrerai, new species, paratype 
(ventral and right lateral aspects). CHILE, Parinacota, “Arica,” Co- 
tacotani, 4500 m, II-28-48, (genitalic dissection # 96118-CDM) in 
CAS. Scale = 1 mm. 


with lateral margins broadly angulate at anterior one-fourth, then 
emarginate to nearly parallel-sided caudal one-third, where 
minutely serrate to caudal tip which exceeds width of pectines, cau- 
dal cleft scarcely exceeding pectines cephalad; pectines not minute, 
length each half usually about twice its breadth, anteriorly and pos- 
teriorly nearly or quite truncate; the tines many, inconspicuous. 
Gnathos in lateral view scarcely sclerotized, slightly exceeding 
pectines caudad, and not very divergent ventrally from uncus. 

Female. Head. As male, antennal shaft about twice length of 
club. Wings. Tawny to very pale fulvous (Figs. 47, 49). Forewing, 
average length 11.9 mm [range 10-15 mm] (n = 35). Dorsal surface 
as male; ventral surface pale fulvous much expanded, fuscous border 
greatly reduced, or faded to nearly obsolete. Genitalia. Eighth ster- 
nite in ventral view sclerotized and broadly crescentic, its greatest 
width caudolaterally, its length about one-fourth to one-third its 
width. Apophysis anterioris in lateral view (Figs. 28, 29) somewhat 
to scarcely produced cephalad of junction with lamella postvaginalis. 
Lamella postvaginalis in ventral view narrowly united, each half pro- 
duced ventro-cephalad forming a slight anterior bulge, not nipple- 
like, medially not produced ventrally. Antrum dorsally sclerotized 
and with a longitudinal fold, ventrally caudal and lateral one-half 
membranous. Ductus bursae well sclerotized, in right lateral view 
ductal constriction not very abrupt, the sinus U-shaped; left lateral 
pocket slightly to prominently produced. 

Type. Mabille’s type, a female, is presumably in the Muséum Na- 
tional Histoire Naturelle, in Paris, France. It was collected in Pert. 


Diagnosis and discussion. The material from Ar- 
gentina I have seen (8 specimens) is all from the pre- 


81 


Fic. 27. Female genitalia of H. tentativa, new species, paratype 
(ventral and right lateral aspects). PERU, Ay[acucho], Rio Apacheta 
4200 m, 13°21’S, 74°39’W, 1-24-99, G. Lamas, (genitalia dissection 
# 26309-CDM), in MUSM. Scale = 1 mm. 


Andean Nevadas del Aconquija, just west of San 
Miguel de Tucuman in provincias Catamarca and Tu- 
cuman. The males (4 specimens) are somewhat vari- 
able in their genitalia (see Fig. 23, # J 6213, from 
Catamarca) and may not represent a single species, 
but I am content to consider them all minor variations 
of H. boulleti until such time as more and better mate- 
rial can be studied. 

Males of H. boulleti have a stigma with a microan- 
droconial mass that is usually neither conspicuous nor 
yellow, they have two pairs of metathoracic tibial 
spurs, and usually lack wing borders above that are 
deeply and broadly cut by fulvous along the veins. This 
species is very pale fulvous above, not orange fulvous; 
and it seems to fly alone, not within the ranges of most 
other species of the boulleti group, save for H. her- 
rerai. Its range does evidently overlap those of the next 
species and of H. peruana slightly in a narrow region 
of Puno, Pert, northwest of Lago Titicaca near the 
border of Departamento Cuzco (Fig. 59). Hylephila 
boulleti seems to be on the wing through most of the 
year except for June through August. 

This species is the most commonly collected mem- 


82 


28 


Fic. 28. Female genitalia of H. boulleti (ventral and right lateral 
aspects). CHILE, El Loa, Geiser el Tatio, 4360 m, I-29-65, J. Her- 
rera, (genitalic dissection # 96218-CDM), in CAS. Scale =1 mm. 


ber of the boulleti group. Indeed, I have seen a total of 
55 males and 40 females (27 males, 23 females dis- 
sected) representing 32 localities in Pert, Chile, Bo- 
livia, and Argentina; but the samples almost always 
consisted of one to three specimens, except for one lo- 
cality in Chile and another in Argentina; where the 
sample was six specimens. It wasn’t until the mid- 
1990's when Arthur M. Shapiro made five collections, 
under a CONAF collection permit issued 18 Novem- 
ber 1994 in Arica, Chile, by Juan Silva, Director, 
CONAF, Ist region, that samples of up to 25 were 
taken. Shapiro 1995, in litt. stated that at one locality 
the H. boulleti “were as common as I have ever seen 
any skipper anywhere; I could have taken hundreds, if 
not thousands.” The skippers were (in his words) “in- 
credibly abundant on monocultures of the apparent 
host (oviposition substrate), Deyeuxia breviaristata 
Wedd., a small wiry grass that grows just (above) the 
margins of boggy and saline environments.” The fa- 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 29. Female genitalia of H. boulleti (ventral and right lateral 
aspects). ARGENTINA, Tucuman, Huaca Huasi, Lago Nostra, 4250 
m, XII-26-78, Halloy, (genitalic dissection # 96214-CDM), in IML. 
Scale = 1 mm. 


vored nectar source was a small composite, Werneria, 
that grows flat on the ground in boggy turf, (fide 
Shapiro). He observed oviposition on Deyeuxia, often 
on plants that seemed dry and desiccated, presumably 
owing to the rains having just started. 

Twenty-three ova were recovered by dissection 
prior to KOH treatment (see MacNeill & Herrera 
1999:279) from three females (# 2° 6218, # 2° 6264, and 
# 2 6265), and these averaged 0.8 mm in diameter and 
0.58 mm in height. The diameter ranged from 0.75 to 
0.83 mm, and the height ranged from 0.55 to 0.65 mm. 
The three females separately had different averages: 
# ° 6218 measured the largest with 9 ova averaging 
0.82 x 0.60 mm, and # 9 6265 measured the smallest 
with 8 ova averaging 0.78 x 0.57 mm. The most com- 
mon (not average) measurement of the 23 ova was 0.8 
x 0.6 mm, a fairly high-domed egg, by and large. The 
reticulation was very weakly evident at 100x magnifi- 
cation. 


Hylephila rossi MacNeill, new species 
(Figs. 3c, 24, 30, 52, 53, 58) 
Description. Male. Head. Vestiture buff tinted fulvous mixed 


with black hairs, eyelash black. Palpi with mostly black third seg- 
ment emergent from buff hairs of second segment. Antennal club 


VOLUME 56, NUMBER 2. 83 


Fic. 30. Female genitalia of H. rossi, new species, paratype (ventral 
and right lateral aspects). PERU, Cuzco-Puno, La Raya, 4318 m, IV-19- 31 
71, J. Herrera, (genitalic dissection # 23829-]H), in CAS. Scale = 1 mm. 


anteriorly buff merging to fulvous and black above, to white below, 
posteriorly black; nudum orange-brown, apiculus width equal to 
length; shaft more than twice club length. Body. Hind legs missing. 
Wings. Narrow, pointed. Forewing length 12.5 mm (n = 1). Stigma 
rather narrow; microandroconial mass brownish gray; apical, middle, 
and lower brush patches present; post-stigmal patch fairly broad. 
Orange-fulvous of apical, and subterminal spots, also of narrow post- 
median macular band (offset from the subterminal spots), and of 
lower one-half of discal cell (expanded end cell), somewhat re- 
stricted by fuscous of costal area, of upper half of discal cell, of space 
Cu2-2A basad of stigma, and of broad, slightly indented border. 
Fringes above fuscous. Below, upper element of stigma pocket di- 
verging from cubitus vein of discal cell about equidistant between 
origins of veins Cul and Cu2, extending nearly or quite basad to 
vein Cu2, lower elements large (Fig. 3c), veins not whitened. Hind- 
wing above with orange-fulvous very restricted to posterior arm of 
postmedian macular band from M1—M3 to Cul—Cu2, and vein 2A 
also fulvous. Below fulvous expanded to base in space Cu2—2A and 
in anterior arm of postmedian macular band across space Rs—M1. 
Veins buff, slightly paler than fulvous ground color, but not 
whitened. Fringes sullied orange, vannally orange. Genitalia. 

Kighth tergite lateral margin broadly concave, not emarginate just 
before eurdal margin; terminal bristle-sockets slightly enlarged be- 
fore caudal margin. Valva (Fig. 24) in lateral view somewhat narrow, 

ventral margin convex, caudally emarginate to beak, basal margin 
straight, dorsal margin slightly concave to caudal beak where knobs 
dorsad of horizontal cleft several; length basal margin about equal to 
one and one-half depth of valva. Penis proximally slightly curved 


Fic. 31. Female genitalia of H. peruana (ventral and right lat- 
eral aspects). PERU, Junin, Pachachaca, 4201 m, IV-9-71, J. Herrera 
(genitalic dissection # 93872-JH), in CAS. Scale = 1 mm. 


dorsad, its length exceeding length of genitalic capsule but not twice 
length of valva; titillators small, nearly symmetric, broadly thorn- 
like, sclerotically strapped to penis; cornuti three, asymmetric, one 
slender, elongate, minutely bidentate, the others small, thorn-like, 
unidentate. Juxta narrow, elongate, with ventrocaudal clefts not 
quite one-half length of juxta, separated median floor not reaching 
caudal margin of juxta. Uncus in dorsal view more or less triangular, 
posteriorly tapered (where minutely serrate) to caudal tip which 
scarcely exceeds width of pectines, caudal cleft scarcely exceeding 
pectines cephalad; pectines small, length each half less than twice its 
breadth, anteriorly and posteriorly nearly truncate; in lateral view 
uncus dorsally arched, pectines conspicuously so. Gnathos in lateral 
view scarcely sclerotized, not divergent ventrally from uncus, and 
not exceeding pectines caudad. 

Female. Head. As male but buff areas paler. Body. Hind tibia 
with upper pair of spurs much reduced, lower pair long. Wings. 
Forewing length 12.5 mm (n = 1). Above, markings rich fulvous, re- 
stricted slightly more than in male, but subterminal spots inclined 
and scarcely offset from postmedian macular band. Hindwing with 
fulvous along veins penetrating fuscous border nearly to termen. Be- 
low, fulvous expanded; veins Rs to Cu2 somewhat whitened. Fringes 
both wings above and below sullied white. Genitalia. Eighth ster- 
nite in ventral view broad, its greatest transverse width cephalad 
about twice its length. Apophysis anterioris in lateral view (Fig. 30) 
produced somewhat cephalad of junction with lamella postvaginalis. 


84 


36 


3/ 


Fics. 32-37. Adults of species of Hylephila, lett side = dorsal 
surface; right side = ventral surface; approximately 1x. 32, H. her- 
rerai, new species holotype 6, same specimen as in Fig. 4a. 33, H. 
herrerai, new species, paratype °, same specimen as in Fig. 26. 34, 
H. pseudoherrerai, new species holotype, same specimen as in Fig. 
4b. 35, H. pseudoherrerai, new species, paratype ¢, same specimen 
as in Fig. 4c. 36, H. pallisteri, new species holotype ¢, same speci- 
men as in Fig. la. 37, H. pallisteri?, 3, same specimen as in Fig. 1b. 


Lamella postvaginalis in ventral view very narrowly united forming a 
distinct V at the junction, from which, in lateral view, each half pro- 
duced but scarcely bulging cephalad. Antrum dorsally sclerotized 
and with longitudinal fold, caudal half membranous ventrally and 
laterally. Ductus bursae sclerotized, left lateral pocket evident in lat- 
eral view but not prominent. 

Types. Holotype 3 PERU, (Puno), 10 mi N. of Ayaviri, III-1-51, 
E. S. Ross & A. E. Michelbacher (genitalic dissection # 6 3821-]H), 
in CAS. Paratype. 1 2 PERU, Cuzco-Puno, La Raya, 4318 m, IV-19- 
71, J. Herrera (genitalic dissection # 293829-JH), in CAS. 

Etymology. This species is named for Edward S. Ross, curator 
emeritus of the California Academy of Sciences, one of the collec- 
tors of the single male known. 


Diagnosis and discussion. Hylephila rossi is 
closely related to H. boulleti, and the two known spec- 
imens are from the only region of overlap known be- 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Zé 


ite 


44 


Fics. 38-45. Adults of species of Hylephila, left side = dorsal 
surfaces; right side = ventral surfaces. Approximately 1x. 38, H. 
blancasi, new species holotype ¢, same specimen as in Fig. le. 39, 
H. blancasi, new species, paratype ¢, PERU, Huanuco, illegible lo- 
cality, Holland collection, (genitalic dissection # 5 6280-CDM), in 
CMNH. 40, H. tentativa, new species holotype d, same specimen as 
in Fig. ld. 41, H. tentativa, new species, paratype °, same specimen 
as in Fig. 27. 42, H. shapiroi, new species holotype ¢, same speci- 
men as in Fig. 2a. 43, H. galera, holotype ¢, same specimen as in 
Fig. 2b. 44, TEL galera? 3, same specimen as in Fig. 2c. 45, H. 


45 


galera? 3, same specimen as in Fig. 2d. 


tween H. boulleti and H. peruana (Figs. 58, 59). This 
species is much darker with much less fulvous above in 
both sexes than H. boulleti, and the fulvous is much 
more orange than the pale fulvous of the latter. Both 


VOLUME 56, NUMBER 2 


Fics. 46-51. Adults of species of Hylephila, left side = dorsal 
surfaces; right side = ventral surfaces. Approximately 1x: 46, H. 
boulleti, 3, same specimen as in Fig. 3a, 47, H. boulleti, 2, same 
specimen as in Fig. 28. 48, H. boulleti, , CHILE, Parinacota, Re- 
serv. Nac. Salar de Surire, no. shore, 4250 m, XI-22-94, A. M. 
Shapiro, in CAS. 49, H. boulleti, 2, same data as specimen in Fig. 48. 
50, H. boulleti 3, same specimen as in Fig. 5d. 51, H. boulleti, 2, 
ARGENTINA, Tucum4én, Las Animas Portezuelo, 4540 m, I-26-79, 
Dominquez, in AMNH. 


the male and female genitalia of each are also slightly 
different (cf. Figs. 22, 24 and Figs. 28, 30). It flies dur- 
ing March and April. 


Hylephila peruana Draudt 
(Figs. 3d, 5e, 8, 11, 25, 31, 54, 55, 56, 57, 59) 


Hylephila lima (sic) Dyar (nec Plétz), 1913. Proc. U.S. 
Nat Mus. 45:639. 

Hylephila peruana Draudt, 1923 [in Seitz] Gross 
Schmet. Erde 5:929, pl. 180 f.; MacNeill & Herrera, 
1999. J. Lepid. Soc. 52:279-280, 291. 

Hylephila boulleti peruana, Evans, 1955. Cat. Amer. 
Hesp. part IV, p. 314; Bridges 1983. Lepid. Hesp. 
notes on species group names, part 1:92; Mielke, 
1993. Rev. Bras. Entomol. 37:622, figs. 93-97; 


Fics. 52-57. Adults of species of Hylephila, left side = dorsal 
surfaces; right side = ventral surfaces. Approximately 1x: 52, H. 
rossi, new species holotype ¢, same specimen as in Fig. 3c. 53, H. 
rossi, new species, paratype °, same specimen as in Fig. 30. 54, H. 
peruana, 6 same specimen as in Fig. 3d. 55, A. peruana °, same 
specimen as in Fig. 31. 56, H. peruana 6, PERU, Cuzco, Cuzco-Ol- 
lantaitambo Rd.,VII-7-84, 3000 m, S. Courtney & P. Stern, in CAS. 
57, H. peruana ? PERU, Junin, vic. Abra Anticona, 4843 + 100 m, 
X-19-83 A. M. Shapiro (genitalic dissection # 9 6143-CDM), in CAS. 


Mielke & Schroeder, 1994. Senckenberg. Biolog. 

73:142, fig. 20. 

Hylephila boulleti perunana, (lapsus) Shapiro, 1985. 
Stud. Neotrop. Fauna and Environ. 20:9,10, figs. 
5(1), 6(1). 

Description. Male. Head. Vestiture buff, tinted orange; eye- 
lash black. Palpi with third segment dorsally black, ventrally buff, 
distinctly emergent from buff vestiture of second segment. Antennal 
shaft long, about thrice length of club; club with nudum pale brown, 
buff proximally, darkening through pale orange to brown at apiculus, 
and nearly one-half length of club. Wings. Somewhat broad but not 
stubby or rounded. Forewing average length 13.6 mm [range 12-14 
mm] (n = 15). Dorsal surface, stigma broad, microandroconial mass 
conspicuous, yellow; apical, middle, and lower black brush patches 
present and united, forming a continuous border around the mi- 
croandroconial mass, poststigmal patch broad and conspicuous. In- 
ner edge fulvous subterminal spots usually offset distad from inner 


86 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


40.-—<—_—<$____—-_————- pees ; UNGUED , 


CENTRAL WESTERN SOUTH AMERICA 
SCALE 
° too 200 «300 400 S00 = anges 


SINUSOIDAL PROJECTION 


Fic. 58. Distribution map for eight species of Hylephila: Solid diamond = H. herrerai, new species; outlined triangles = H. pseudoherrerai, 
new species; outlined circles = H. pallisteri, new species; outlined squares = H. blancasi, new species; outlined diamond = H. tentativa, new 
species; solid circle = H. shapiroi, new species; solid inverted triangles = H. galera; solid squares = H. rossi, new species. 


VOLUME 56, NUMBER 2 


CENTRAL WESTERN SOUTH AMERICA 


SCALE 
° 100 200 300 aco sco G00 MALES 
o 200 400 600 600 KILOMETERS 


SINUSOIDAL PROJECTION 


Fic. 59. Distribution map for two species of Hylephila: Outlined circles = H. boulleti; solid diamonds = H. peruana. 


88 


edge of postmedian macular band. Fulvous broadly warm-orange 
costad and basad in discal cell, usually extending well into fuscous 
border along veins, but occasionally border broad and scarcely in- 
dented (cf. Figs. 54 and 56). Fringes orange-fulvous. Below, stigma 
pockets broad; upper element spanning distance between origins of 
veins Cul and Cu2, diverging from discal cell cubitus vein much 
nearer to vein Cu2 than to vein Cul (Fig. 3d). Fulvous more exten- 
sive than above, veins R5 to M3 distally whitened. Fringe basally 
narrowly whitish, then fuscous, then terminally sullied fulvous. 
Hindwing above orange-fulvous as on forewing; vertex of chevron- 
shaped postmedian macular band in space M1—MB3 usually conspic- 
uously extended basad well into discal cell, and anterior arm present 
in space Rs-M1. Fringe orange fulvous. Below, fulvous paler than 
above; veins Rs to M3 whitened but not conspicuously so to naked 
eye; often a trace of an additional postmedian, elongate, black dash 
in upper part of space M1—M3. Fringe as on forewing. Genitalia. 
Eighth tergite lateral margin broadly concave mid-length but not 
distinctly emarginate just before caudal margin (Fig. 5e); terminal 
bristle-sockets enlarged and rounded just before caudal margin (Fig. 
8). Valva (Fig. 25) in lateral view somewhat broad, ventral margin 
proximally convex, basal margin scarcely convex, dorsal and ventral 
margins caudally concave to caudal beak where knobs dorsad of hor- 
izontal cleft numerous; length basal margin greater than one and 
one-half times depth of valva. Penis proximally nearly straight or 
slightly curved dorsad, its length slightly exceeding length of geni- 
talic capsule and nearly twice length of valva; titillators asymmetric, 
the left large, laterally compressed, thorn-like with a very broad 
base, the right much reduced, thorn-like, both sclerotically strapped 
to penis; cornuti asymmetric, one narrow, basally elongate, and 
minutely bidentate, the other basally broad and thorn-like, biden- 
tate. Juxta with ventrocaudal clefts long, about one-half length of 
juxta; separated median floor reaching caudal margin of juxta. Uncus 
in dorsal view anteriorly rounded but posteriorly tapered (where 
minutely serrate) to broad caudal tip that greatly exceeds width of 
pectines, caudal cleft not exceeding pectines cephalad; pectines 
minute, length each half medially more than twice its breadth, dis- 
tinctly tapered mesad anteriorly; the tines conspicuous, few; in lat- 
eral view uncus and pectines dorsally nearly flat. Gnathos in lateral 
view scarcely sclerotized, not divergent ventrally from uncus, 
scarcely exceeding pectines caudad. 

Female. Head. As male but antennal shaft slightly shorter not 
quite thrice length of club. Wings. Orange to tawny fulvous above 
somewhat more restricted than in male owing to slightly broader 
fuscous border. Forewing average length 14 mm [range 13-15 mm] 
(n = 15). Below as male, but forewing Border usually greatly reduced 
in spaces M1-M2, M2-M3, and Cu2-2A. Genitalia. Eighth ster- 
nite in ventral view sclerotized, broad, more or less quadrangular 
(rounded ventrad in lateral view), with anterior margin straight or 
emarginate, its length in ventral view nearly or quite one-half its 
width. Apophysis anterioris somewhat produced cephalad of junc- 
tion with lamella postvaginalis (Fig. 31). Lamella postvaginalis in 
ventral view narrowly united, each half produced ventrocephalad 
forming a slight anterior bulge, not nipple-like, medially not pro- 
duced ventrally. Antrum dorsally sclerotized, with longitudinal fold, 
anterolaterally with increased sclerotization just before ductal con- 
striction, ventrally membranous. Ductus bursae well sclerotized, in 
lateral view massive just beyond deep V-like ductal constriction, in 
ventral view left lateral pocket slightly produced. 

Types. Lectotype ¢ (designated by Mielke 1993:622) in Sencken- 
berg Museum, Frankfurt, Germany [SMF L 4068 (Genit. Prap. O. 
Mielke #532)]. PERU, Cuzco, 4500 m Fig. in Seitz (1923: Taf. 180 £ 
underside). Paralectotypes. Two specimens (designated by Mielke 
1993:622) in Senckenberg Museum, Frankfurt, Germany. 1 2 SMF L 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


4069, Fig. in Seitz (1923: Taf.180 f); PERU, Cuzco, 3500 m. 1d SMF 
L 4070, Fig. in Seitz (1923: Taf. 180 f). PERU, Cuzco, 4500 m. 


Diagnosis and discussion. In the boulleti group of 
Hylephila, males of this species are arguably the speci- 
mens most easily identified without dissection, because 
of their conspicuous stigma with its yellow microandro- 
conial mass, their extremely abbreviated uncal pectines 
(which can often be seen by merely gently brushing 
the abdominal tip from above), and their distribution 
in Pert: north and northwest of the department of 
Puno, where H. peruana occurs with several other 
members of the group (see maps, Figs. 58, 59). 

This is the member of the boulleti group most com- 
monly collected in Pert. I have seen 22 males and 20 
females (18 males, 17 females dissected) from 18 lo- 
calities in Pert. Flight records suggest perhaps four 
broods through the year with gaps appearing during 
March, June, September, and November—December. 

Nine ova were dissected from three females (# 2 
6377, #2 6378, # 2 6379) prior to KOH treatment. Two 
eggs from # 2 6377 measured an average 0.75 x 0.63 
mm, and also a pair of eggs from # ° 6378 averaged 
0.85 x 0.55 mm. Five ova from # 2 6379 averaged 0.83 
x 0.57 mm, and the average of all nine ova is 0.81 x 
0.57 mm. The most common measurement for all the 
eggs was 0.85 x 0.55 mm. The eggs were all very simi- 
lar to eggs of H. boulleti including the very weak retic- 
ulation at 100 x magnification, although the ova of this 
species proportionately do not seem to be as highly 
domed as are those of H. boulleti. 


ACKNOWLEDGMENTS 


I am grateful to the following individuals and institutions for pro- 
viding me with loans of material or information: Lee D. Miller and 
the Allyn Museum of Entomology (AME), Florida Museum of Nat- 
ural History, Sarasota, Florida; Fred H. Rindge and The American 
Museum of Natural History (AMNH), New York, New York; Philip 
R. Ackery and the Natural History Museum (BMNH), London, 
England; Paul H. Arnaud Jr. and the California Academy of Sciences 
(CAS), San Francisco, California; John E. Rawlins and the Carnegie 
Museum of Natural History (CMNH), Pittsburgh, Pennsylvania; 
Rauil Cortez and the Instituto de Entomologia, Universidad Metro- 
politania de Ciencias de la Educacién (IEUM), Santiago, Chile; the 
late Abraham Willink and the Instituto de Zoologia, Fundacion 
Miguel Lillo (IML), San Miguel de Tucumn, Argentina; Zsolt Balint 
and the Hungarian Natural History Museum (HNHM), Budapest, 
Hungary; Julian P. Donahue and the Natural History Museum of 
Los Angeles County (LACM), Los Angeles, California; Gerardo 
Lamas and the Museo Nacional de Historia Natural, Universidad 
Mayor de San Marcos (MUSM), Lima, Pert; Arthur M Shapiro and 
the Bohart Museum of Entomology, University of California (UCD), 
Davis, California; Olaf H. H. Mielke and the Departamento de Zo- 
ologia, Universidade Federal do Parana (UFPC), Curitiba, Brasil; 
John M. Burns and the National Museum of Natural History, Smith- 
sonian Institution (USNM), Washington, D.C.; and the late Luis E. 
Pefia, Santiago, Chile. I am indebted also to Yu-Feng Hsu of the Na- 
tional Taiwan Normal University, Taipei, Taiwan who hand-carried 
several of Evans's types to me from the BMNH. Genitalic drawings 
were expertly detailed by Shannon Bickford, in Fresno, California, 


VOLUME 56, NUMBER 2 


and scanning electron micrographs were produced by Darrell Ubick 
at the CAS. Arthur Shapiro continued his tireless collecting efforts 
and observations in South America, and I particularly want to thank 
Juan Silva of Arica, Chile, Director of CONAF, First Region, for 
granting A. M. Shapiro the collecting permit that made his efforts in 
Chile possible. The artwork was partially funded by a CAS In-House 
Research Fund FY 1994-95 grant, and some of the manuscript and 
publication costs were covered by CAS Entomology Research 
Funds. Julie Parinas, Paul Amaud, and Vincent Lee again repeat- 
edly were tolerant and helpful in solving my many entreaties for as- 
sistance with my “new” word processor. John Burns and George 
Austin very kindly read the manuscript and offered numerous sug- 
gestions which much improved the paper, for which I am most 
grateful. My thanks to them all. 


LITERATURE CITED 


BILLBERG, G. T. 1820. Enumeratio insectorum in museo Gust. Joh. 
Billberg (Holimiae) Gadel. i-iv + 1138 pp. 

BRIDGES, C. A. 1983. Lepidoptera: Hesperiidae, Notes on species- 
group names. Pp. I.1—1.129, I.1—II.41, I1.1—I11.62, IV.1-IV.30, 
V.1-V.13. C.A. Bridges, Urbana, Illinois. 

Draut, M. 1923. B. Grypocera, breitképfige Tagfalter. In Seitz, A. 
(ed.), Die Gross-Schmetterlinge der Erde. 5. Die americanis- 
chen Tagfalter. Pp. 836-1022, pls. 189-208. Alfred Kemen, 
Stuttgart. 

Dyar, H. G. 1913. Results of the Yale Peruvian expedition of 1911. 
Lepidoptera. Proc. U.S. Natl. Mus. 45:622-649. 

Evans, W. H. 1955. A catalogue of the American Hesperiidae, 
indicating the classification and nomenclature adopted in the 
British Museum (Natural History). Part IV. Hesperiinae and 
Megathyminae. British Museum, London. 499 pp., pls. 
54-88. 


89 


HaywarD, K. J. 1973. Catalogo de los Ropaloceros [sic] Argentinos. 
Opera Lilloana 23:318 pp. 

Lewis, G. 1973. Butterflies of the World. Chicago. Pp. (vi)—xiy, 
209-312, pls. 1-208. 

MABILLE, P. 1906. Description d'une Hesperide (Lep.). Bull. Soc. 
Entomol. France, no. 6:67-68. 

MACNEILL, C. D. & J. HERRERA G. 1999. Studies in the genus 
Hylephila Billberg, I. Introduction and the ignorans and 
venusta species groups (Hesperiidae: Hesperiinae). J. Lepid. 
Soc. 52:277-317. 

MIELKE, O. H. H. 1993. Sobre os tipos de Hesperiidae (Lepi- 
doptera) neotropicais descritos por M. Draudt. Rey. Bras. En- 
tomol. 37:611-638. 

MIELKE, O. H. H. & H. G. SCHROEDER. 1994. Insecta: Lepi- 
doptera: Hesperiidae von M. Draudt aus der neotropis 
beschrieben arten. Die typen and typoide des natur-museums 
Senckenberg Nr. 82. Senckenbergiana Biologica 73:135-158. 

PENA G., L. E. & A. J. UGARTE P. 1997. Las mariposas de Chile, the 
butterflies of Chile. Editorial Universitaria, Santiago. [9] + 357 
pp. figs. (on p. 227). 

PLOTZ, C. 1883. Die Hesperiinen-gattung Hesperia auct, und ihre 
arten. Stet. Ent. Zeit. 44:195-233. 

Sapiro, A. M. 1985. Behavioral and ecological observations of Pe- 
ruvian high-Andean Pierid butterflies (Lepidoptera). Stud. on 
Neotrop. Fauna and Environ. 20:1—133, Figs. 1-6 

. 1993 [1991]. Convergent evolution in western North 

American and Patagonian skippers (Hesperiidae). J. Res. Lepid. 

30:162-174, 4 figs. 

. 1994 [1992]. Why are there so few butterflies in the high 

Andes? J. Res. Lepid. 31:35-56, pls. I-II. 


Received for publication 5 April 2001; revised and accepted 7 No- 
vember 2001. 


Journal of the Lepidopterists’ Society 
56(2), 2002, 90-97 


DEW-DRINKING BY MALE MONARCH BUTTERFLIES, DANAUS PLEXIPPUS (L.) 


DENNIS FREY, ROBERT ROMAN AND LINDSAY MESSETT 
Biological Sciences Department, California Polytechnic State University, San Luis Obispo, Califomia 93407, USA 


ABSTRACT. We examined the early moming activity of overwintering monarch butterflies at a central coast California site during their 
period of “mass-mating” from mid-January to mid-March in 1998 and 1999. The first year of the study took place during El Nifio weather when 
regional precipitation was approximately 50% greater than normal. The second year occurred during La Nifia conditions with approximately 
15% less precipitation than normal. On “rain-free” mornings many of the butterflies left aggregations and flew to an adjacent meadow where 
they landed on grass and drank from dew droplets. Males were much more likely to make morning meadow visits than females and such males 
had 6.7% higher body moisture than males that remained in the clusters. The rate of male meadow use increased seasonally both years and 
males were frequently seen attempting to mate following dew-drinking meadow visits. Early moming meadow-males resembled males at- 
tempting to mate in the morning more than males that were not mating, i.e., that remained in aggregations. Meadow-males were smaller, had 
relatively smaller dry-weight abdomen mass, and had more wing damage in addition to their greater moisture content. Variation in local solar 
radiation, wind speed and evapotranspiration during the previous 24 h was positively associated with early momming meadow use by both males 
and females. Male mating effort during the previous day was positively associated with early morning dew-drinking by males but not females. 
Three hypotheses regarding dew-drinking activity are considered. The results are consistent with two perspectives: dew drinking results from 
dehydrating activity such as courtship and spermatophore transfer during the previous day and/or it represents strategic male behavior in an- 


ticipation of “need” for future reproductive effort. 


Additional key words: Danaus plexippus, dew-drinking, mating effort, dehydration, E] Nifio, La Nifa. 


Each fall monarch butterflies, Danaus plexippus 
(L.), migrate from an extensive late summer breeding 
range in North America to geographically restricted 
overwintering habitats (Brower & Malcolm 1991). 
Monarchs from eastern breeding populations overwin- 
ter in several mountainous areas dominated by Oya- 
mel fir forests near Mexico City (Urquhart & Urquhart 
1978, Brower 1985, Calvert & Brower 1986). Those 
originating west of the Rocky Mountains overwinter at 
wooded sites located along the Pacific Ocean between 
Bolinas, California and Enseneda, Mexico (Sakai & 
Calvert 1991, Lane 1993). 

The general eco-region macroclimate where these 
sites are located, along with local topography and veg- 
etation, provide microclimate conditions favorable for 
monarch overwintering (Brower et al. 1977, Calvert et 
al. 1982). Conditions thought to be important for over- 
wintering include: (1) generally cool temperatures that 
minimize the rate of butterfly activity and rate of body 
fat utilization (Chaplin & Wells 1982, Masters et al. 
1988, Alonso-Mejia et al. 1997), (2) a pattern of daily 
minimum temperature that reduces the likelihood of 
lethal freezing (Alonso-Mejia et al. 1997), (3) enough 
moisture to minimize desiccation (Calvert & Lawton 
1993), (4) low wind velocity to minimize disruption 
and desiccation (Leong et al. 1991), and (5) exposure of 
butterflies in cluster formations to brief mosaics of di- 
rect solar radiation to enable periodic thermoregula- 
tory basking (Calvert & Brower 1986, Frey et al. 1992). 

Overwintering at Mexico sites occurs during the re- 
gional dry season but dehydrating conditions are re- 
duced somewhat because these sites occur at high ele- 
vations (2400 to 3600 m) where cloud forests provide 
relatively high humidity and closed-canopy forests re- 
sult in wind abatement (Calvert & Brower 1986). 


Overwintering at California sites on the other hand oc- 
curs during the wet season, yet desiccation is a prob- 
lem for monarchs. Moisture-producing storms de- 
crease in frequency during February and March, 
occur sporadically, last for only brief periods and are 
often followed by many days of dry conditions. For ex- 
ample, during the wet 1998 EI Nifio event, on 58 days 
in January, February and March, moisture loss 
through evapotranspiration exceeded moisture gain 
from precipitation along the central coast of California 
(data source: California Department of Water Re- 
sources, California Irrigation Management Informa- 
tion System from weather stations located in 
Guadalupe, California). This potential for desiccation 
was even more pronounced during the dry 1999 La 
Nina winter. 

An intense period of mating occurs between late 
January and early March at both Mexico and Califor- 
nia overwintering sites during which males compete 
vigorously for mating opportunities (Hill et al. 1976, 
Tuskes & Brower 1978, Van Hook 1996, Frey et al. 
1998). During a lengthy copulation the male transfers 
a spermatophore to his partner's reproductive tract, 
which may equal 10% of his body weight (Oberhauser 
1988). Monarch spermatophores contain a small am- 
poule of spermatozoa along with various nutrients, but 
consists primarily of water (up to 93%) and females 
may use this “transferred” water for somatic mainte- 
nance (Oberhauser 1992, Pham 1997). At California 
overwintering sites mating could increase water deficit 
for males due to the high water content of sper- 
matophores and the seasonal increase in evapotranspi- 
ration, solar radiation and temperature. Overwintering 
sites in Mexico and California are usually associated 
with a nearby source of water (Calvert & Brower 1986, 


VOLUME 56, NUMBER 2 


Leong 1990, Leong et al. 1991, Bell et al. 1993), but 
cause and effect relations of monarch hydration ac- 
tivity are not fully understood and hydration behavior 
has not been studied relative to male mating strategy. 

At central coast sites in California monarchs remain 
relatively inactive throughout much of the overwinter- 
ing period and spend most of the time in cluster for- 
mations (Hill et al. 1976, Tuskes & Brower 1978, Frey 
& Leong 1993, Frey et al. 1998). On relatively clear 
days when temperatures exceed the flight threshold 
many butterflies leave the aggregations and engage in 
a variety of activities. The majority of them land on 
sun-exposed canopy vegetation, while some nectar on 
the limited nearby flowering plants. At the North 
Beach Campground overwintering site located in 
Pismo Beach, California many of the butterflies that 
emerge from aggregations in the early morning fly to 
an adjacent meadow where they land on dew covered 
grass and appear to drink from dew droplets (pers. 
obs.). Small meadows regularly occur adjacent to west- 
ern North America winter habitat or within the small 
gaps found in the stand of trees comprising the habitat. 

In this study we examined the early morning activity 
of monarch butterflies at a central coast California 
overwintering site during the mass mating phase (Jan- 
uary through March) of 1998 and 1999. Our objectives 
were to (1) describe the departure pattern of males 
and females as they left aggregations to visit a nearby 
meadow, (2) compare moisture content and morphol- 
ogy of males that visited the meadow to those that re- 
mained in aggregations or were engaged in other ac- 
tivities, (3) test whether weather variables from either 
the current day or from the previous day could explain 
any patterns from objectives 1 and 2 above, and (4) 
contrast meadow visiting activity and moisture content 
during two seasons differing greatly in regional mois- 
ture regimes. 


MATERIALS AND METHODS 


The overwintering site. This study was carried 
out in the southeast corner of North Beach Camp- 
ground, Pismo Beach State Park, Pismo Beach, Cali- 
fornia. Throughout the study monarchs formed over- 
wintering aggregations in a stand of trees dominated 
by blue gum eucalyptus (Eucalyptus globulus Labill.) 
along with scattered Monterey cypress (Cupressus 
macrocarpa Hartw.) and Monterey pine (Pinus radiata 
Don). A meadow 60 x 90 m, bordered on the west and 
north by trees, occurs in the southeast quarter of the 
site and is comprised primarily of rip-gut brome (Bro- 
mus diandrus Roth) and veldt grass (Ehrharta ca- 
lycina Smith). The old-growth grass in the meadow 
was cut back to short stubble in early October during 


91 


both years. By the end of March 1998, the grass was 30 
to 40 cm tall and uniformly covered most of the 
meadow. At the end of the overwintering period in the 
second year, i.e., March 1999, the grass was 20 to 30 
cm tall. 

Counts of monarchs in the meadow. Four uni- 
formly spaced east-west transects were established 
across the meadow covering a total length of 240 m. 
On 17 days between 21 January 1998 and 10 March 
1998 we visited the site in the early morning before 
ambient temperature exceeded the flight threshold 
(10-16°C; Masters et al. 1988, Alonso-Mejia et al. 
1993). In 1999 we made 29 morning counts between 
28 January and 26 March. On each of these days dew 
was present on the grass and other meadow vegeta- 
tion. We slowly walked along each transect and 
counted the number of male and female monarchs lo- 
cated within 3 m on either side. A second count was 
done approximately 45 min after the first monarchs 
began flying out from the cluster trees and landing in 
the meadow. 

Macro and micro weather conditions. Pismo 
Beach is located in a Mediterranean type ecoregion 
(Bailey 1978) and the study was conducted during 
what is usually the transition from the wet season to 
the dry season. The first year of the study occurred 
during El] Nifio weather (wetter than average) while 
the second season was considered a La Nifia period 
(drier than average). For each day of the study we 
downloaded weather data from nearby California Irri- 
gation Management Information System (CIMIS) 
recording stations located in Guadalupe and San Luis 
Obispo, California. The Pismo Beach overwintering 
site is located between these stations approximately 19 
km from each station. The precipitation for January 
through March in 1998 and 1999 was approximately 
50% greater (1998) and 15% less (1999) than the 
regional 30-year rainfall average. We restricted our 
observations to rain-free mornings because monarchs 
do not fly during overcast rainy periods. Year-to-year 
differences for all other weather variables on days of 
data collection were not as pronounced as the differ- 
ences in the overall regional precipitation patterns re- 
ported above. In fact, evapotranspiration, solar radia- 
tion, and average wind speed recorded at the nearby 
CIMIS weather stations on days before morning 
meadow counts did not differ significantly between 
years (Table 1). We also recorded on-site air tempera- 
ture in the meadow each day at the time when we ob- 
served the initial fight of a monarch from the cluster 
trees to the meadow and at the time of both of the 
counts of butterflies along the meadow transect. Initial 
flight temperature did not differ between years but 


92 


TABLE 1. Regional weather variables recorded during 1998 and 
1999 on (a) the morning of meadow counts and (b) the day preced- 
ing meadow counts. Values are daily means + SE (standard error). 


Variable! 1998 1999 Pp? 


(a) Temperature of initial fight 11.2+0.4 10.4+0.3 0.106 
(°C) 

Change in temperature 22+04 44+04 <0.001 
(Ge)? 

(b) Evapotranspiration (mm) 19+03 22+0.2 0.459 

Solar radiation (W m7) 144+17 162+10 0.649 

Wind velocity (ms) 27402 25+01 0.344 


' Evapotranspiration, solar radiation, and wind velocity values 
were down-loaded from CIMIS data bases for stations #52 in San 
Luis Obispo, California and #120 in Guadalupe, California main- 
tained by the California Department of Water Resources. 

P-values are from Mann-Whitney tests. 

’ Change in temperature was recorded daily on-site as the differ- 
ence in temperature of the initial morning flight from cluster trees 
versus the temperature during the second morning transect count 
approximately 45 minutes later. 


mornings during 1999 warmed more quickly between 
transect counts than during 1998 (Table 1). 

Moisture content and condition of monarchs. 
Males were captured weekly during 1998 from three 
different categories: (1) dew drinkers: males located in 
the meadow with their probosces extended into dew 
droplets on grass, (2) mating pairs: males that had just 
captured and coupled with a female near the meadow, 
(3) aggregating individuals: males captured with a net 
attached to a long-reach pole from the aggregations 
one hour after ambient temperature exceeded flight 
threshold. Males from the meadow and those mating 
were captured immediately following the second tran- 
sect count of monarchs in the meadow. During 1999, 
males were captured in the meadow as described for 
category 1 above on the mornings of 23 February and 
2 March. Males were captured from clusters as de- 
scribed for category 3 above on 20 February and 27 
February 1999. Mating males were not sampled dur- 
ing 1999. 

The number of wings that had membrane tears or 
portions missing were counted for each butterfly. We 
placed each male individually in a freezer proof zip- 
lock bag and stored them in an ice-chest until they 
were transported to our lab where they were frozen at 
-20°C. These specimens were later weighed (wet 
weight) and then dehydrated in a drying oven at 60°C 
for 40 h. Individuals were weighed immediately on re- 
moval from the freezer to minimize bias due to accu- 
mulation of water that condenses on thawing speci- 
mens. Following dehydration their dried body weight 
was recorded and moisture content was estimated as 
the difference between wet and dry weight. Following 
dry weight measurement, the abdomen was severed 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


from the rest of the body and weighed separately. Dry 
weight of the abdomen relative to total dry body 
weight was used as an index of body condition or an 
approximate “fat-content” index. 

Estimation of male mating effort. We estimated 
male mating effort in two ways. In the first technique 
we videotaped courtship responses of males to pinned 
monarch specimens positioned on platforms in areas 
of high mating activity. From late morning recording 
sessions (55 min duration) we summed the number of 
“visits” by males to the models as well as the number 
of times males landed on them and tried to couple 
with the models. For more details of this approach see 
Falco (1999). These values were converted to standard 
normal scores and used as an index of population-level 
male mating effort. The second technique involved 
late morning counts (approximately 1 h duration) of 
“ground pairs”, i.e., a male attempting to couple with a 
female, in the area where mating attempts frequently 
occurred. We followed the protocol of Frey (1999) and 
converted the number of observed attempts per 
minute to standard normal scores. 


RESULTS 


Entry rates into the meadow. A few monarchs 
were occasionally found in the meadow during count 
#1 before the beginning of morning flight activity. 
They were usually wet with dew, much more so than 
ones collected later in the day, and females were often 
found near males. We used an abdominal palpation 
technique (Van Hook 1999) and found that most of the 
females had a large detectable spermatophore present. 
This suggested that they had mated recently and pos- 
sibly had spent part of the previous night in copula in 
the meadow rather than more typically being attached 
to males that perched in the canopy of nearby trees. 

During the first year of the study the Pismo Beach 
monarch population declined from a peak abundance 
of 125,000 butterflies in late December 1997 to less 
than 1000 individuals by March 1998. During the sec- 
ond year the population declined from 100,000 mon- 
archs in December 1998 to less than 1000 butterflies 
in March 1999. In both years the sex ratio became in- 
creasingly male biased seasonally which is a pattern 
previously reported for this site (Frey & Leong 1993, 
1995, Frey et al. 1998). We computed an index of sex- 
specific increase in meadow abundance to adjust for 
the following factors: (1) differences in abundance and 
sex ratio pattern between years, (2) seasonal declines 
in abundance due to dispersal that began in January 
each year, and (3) slight day-to-day differences in the 
time interval between transect counts. This index was 
computed by dividing the number of new butterflies 


VOLUME 56, NUMBER 2 


TABLE 2. Number of male and female monarchs counted along 
transects in a meadow adjacent to overwintering cluster-trees, sex 
ratios, and per capita meadow entry rate for each sex. Results in- 
clude 17 and 29 sampling dates for 1998 and 1999 respectively. 


Meadow 
Year Sex First count Second count entry rate* 
1998 Male 12.6 + 2.4 66.1 + 12.3 0.031 * + 0.006 
(67%) (85%) 
Female 49+1.1 10.6 + 1.9 0.007" + 0.004 
(33%) (15%) 
1999 Male 2.0 + 0.6 24.3423 0.023 + 0.003 
(65%) (86%) 
Female 10+0.3 41+0.7 0.011 > + 0.003 
(35%) (14%) 


* Individuals entering the meadow per min per 1000 males or fe- 
males remaining in the overwintering area; values with the same let- 
ter subscript do not differ significantly (P > 0.05) by unpaired Mann- 
Whitney U-tests. Counts are means + SE. 


arriving along the transects (i.e., second transect count 
minus initial count) by the time between counts and 
then dividing this value by the number of either males 
or females estimated to be present at the overwinter- 
ing site on a particular date. Population abundance for 
each sex were available from MRR Jolly-Seber census 
estimates made weekly throughout both years of the 
study (DF unpublished data). 

Meadow use rate adjusted for the factors above (i.e., 
per capita meadow entry) did not differ significantly 
between years for either sex (Table 2: males, P = 0.47; 
females, P = 0.16), but male rate was significantly 
greater than female rate for both years (Table 2: 1998, 
P < 0.01; 1999, P < 0.01). In 1998 male per capita en- 
try rate into the meadow, averaged over the season, 
was 4.4 times greater than female rate and 2.1 times 
greater in 1999; by mid-morning the meadow was 
dominated by males. 

Body moisture, morphology and wing condi- 
tion of males. Meadow-captured males had greater 
moisture content than aggregating males during both 
years (Table 3: P < 0.01) but year-to-year differences in 
moisture content were not significant (Table 3: P = 
0.79). During 1998, moisture content for both males 
collected from the meadow and those captured while 
mating early in the morning were significantly greater 
than cluster captured males (l-factor ANOVA & 
Fisher PLSD test, F = 13.57, df = 2, 135, P < 0.01). 
Males captured after spending less than | h in the 
meadow had approximately 6.7% greater moisture 
content than their counterparts collected from aggre- 
gations. 

Males captured from dew-covered grass had signifi- 
cantly less dry weight body mass than cluster-captured 
males during both years (Table 3: P < 0.01). However, 
a significant two-way interaction effect suggested that 


93 


TABLE 3. Moisture content, total dry body mass, and relative 
abdomen mass of male monarchs. Letters that differ within columns 
indicate significant differences (P < 0.05) by pairwise Mann- 
Whitney tests within years. 


Mean Mean 


dry dry 

Mean body abdomen 
Capure Sample moisture mass mass 
Year category size content (%) (mg) (%) 
1998 Meadow 4] 604 1984 284 
Aggregations 47 56° 943> 30» 
Mating 50. 59 # 1982 298 
1999 Meadow 40 614 184° 35° 
Aggregations 4] 57> 9534 434 


the difference between dew-drinking vs. clustering 
male dry mass was more pronounced during 1999 
(Table 3: year x category, F = 11.49, df = 1, 165, P < 
0.01). Meadow-males had 18.5% and 27.3% less dry 
mass than their clustering counterparts in 1998 and 
1999 respectively. 

Difference in relative abdomen mass (i.e., dry ab- 
domen mass divided by dry body mass) among the 
three categories was marginally significant for 1998 
(Kruskal-Wallis H = 5.6, P = 0.059) but field captured 
males had significantly smaller abdomens than cluster- 
captured males (Table 3: P = 0.019). Meadow-males 
had significantly smaller abdomens than cluster-males 
in 1999 (Table 3: P < 0.01). 

The dew drinking and mating males also had more 
damaged wings than males taken from clusters during 
the first year of the study (Two way contingency analy- 
sis; 3 groups X 5 damage categories; X* = 19.6; df = 8; 
P = 0.012). Wing damage pattern was not surveyed 
during year 2. 

Factors affecting meadow use. Neither the vari- 
ation in the temperature of initial morning flights nor 
the variation in the increase in morning temperatures 
were significantly associated with meadow entry rates 
for either sex (Table 4). On the other hand, regional 
solar radiation received during the previous day was 
significantly associated with meadow entry rate for 
both sexes (Table 4: males, P < 0.01; females, P < 
0.05). Evapotranspiration and wind velocity were mar- 
ginal predictors of male meadow use but not female 
use. Days with dehydrating conditions (high solar radi- 
ation, evapotranspiration, and wind velocity) were fol- 
lowed the next morning by high meadow entry rate. 

“Mating effort”, i.e., the standard normal scores of 
the methods outlined above, was significantly as- 
sociated with male per capita meadow-use or dew- 
drinking recorded the following morning (Spearman Z 
= 1.95; N = 20, P = 0.05). When mating activity was 
high, early morning use of the meadow was high on 


94 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 4. Results of Spearman correlation tests of weather variables as predictors of meadow entry rate during the 46 sampling dates of the 
1998 and 1999 monarch mating seasons combined; (a) tests based on temperatures recorded on-site during the morning of meadow counts as 
predictor variables, (b) tests based on weather conditions during the day preceding meadow counts as predictors. 


Influence on male meadow entry rate Influence on female meadow entry rate 
Predictor variable Spearman “x” P Spearman “x” P 
(a) Temperature of initial flights (°C) 1.47 0.142 —0.87 0.386 
Change in temperature during morning (°C) 1.58 0.114 1.66 0.097 
(b) Evapotranspiration (mm) 1.92 0.055 1.41 0.159 
Solar radiation (W m~) 2.85 0.004 2.34 0.019 
Wind velocity (m s“) 1.74 0.082 0.561 0.576 


the following day and vice versa. Female meadow-use 
was unrelated to the previous day’ mating effort 
(Spearman Z = 0.19; N = 20, P = 0.85). 

Female per capita use of the meadow did not 
change significantly during the course of the season 
(Spearman correlation; Z = 1.36, P = 0.18, N = 46). On 
the other hand, male per capita entry into the meadow 
increased significantly as the mass mating season pro- 
gressed (Spearman correlation; Z = 2.80, P = 0.005, N 
= 46). Relatively greater proportions of the male popu- 
lation made early morning visits to the meadow later in 
the season. 


DISCUSSION 


Male and female monarchs flew from overnight 
roosting aggregations to a nearby meadow as morning 
temperature warmed above the flight threshold. Indi- 
viduals of both sexes were observed drinking from dew 
droplets on grass blades, i.e., they had their probosces 
extended into the droplets for long periods of time. 
Males collected from the meadow had 6.7% greater 
moisture content than males collected from aggrega- 
tions, which also suggests that they had been drinking. 
Leong et al. (1992) reported monarch water loss be- 
tween 6% and 10% per day when held without water 
under lab conditions (19.1°C and 44.9% relative hu- 
midity). Males were much more likely to visit the 
meadow than females during the early morning when 
dew was available (Table 2). 

Dew-drinking patterns were similar between the 
two years of this study even though the amount of pre- 
cipitation differed greatly (Table 2). This pattern indi- 
cates that monarch moisture levels are closely regu- 
lated and influenced more by short-term events (e.g., 
the past 24 to 48 h) than over a longer term. In con- 
trast, year-to-year differences in relative abdomen 
mass (Table 3, larger abdomens in 1999) probably re- 
flected overall differences in temperature regimes be- 
tween El Nifio and La Nifia seasons and the fact that 
fat reserves are used more rapidly under higher tem- 
peratures (Chaplin & Wells 1982). Average daily tem- 


perature during January, February, and March of 1998 
approximated the 30-average for the Pismo Beach area 
but were 4%, 3%, and 9% below normal respectively 
during 1999. 

There are several reasons why water may be partic- 
ularly important to males during the mass mating pe- 
riod at overwintering sites beyond a general need to 
reduce negative systemic physiological effects that 
may accompany desiccation. Three hypotheses re- 
garding early morning dew-drinking are given in Table 
5 along with specific testable predictions for each 
hypothesis. Early morning temperatures could influ- 
ence flights from cluster trees (Table 5, Current 
Weather—hypothesis 1) because flight thresholds can 
vary slightly among individuals in overwintering popu- 
lations (Masters et al. 1988, Anson Lui pers. com.). Al- 
ternatively, or in addition, weather associated with de- 
hydration during the previous day could influence 
dew-drinking on the following morning (Table 5, Pre- 
vious Day Weather—hypothesis 2). To test these first 
two hypotheses we combined data from both years of 
the study for the following three reasons: (1) per capita 
meadow entry rate did not differ between years for ei- 
ther sex (Table 2), (2) on-site temperature during the 
initial morning flights did not differ between years, 
and (3) the regional evapotranspiration, solar radia- 
tion, and wind velocity on days preceding our observa- 
tions of dew-drinking did not differ between the two 
years (Table 1). 

Neither temperature-related prediction of the “cur- 
rent weather constraint” hypothesis, i.e., Table 5—hy- 
pothesis 1, was borne out in this study (Table 2 & 
Table 4a). Early morning conditions, however, can in- 
fluence meadow use since monarchs are constrained 
from flying at temperatures below approximately 10°C 
and they seldom fly under overcast conditions (pers. 
obs.). One of the two predictions of the “previous day 
weather” hypothesis, i.e., Table 5—hypothesis 2, was 
confirmed. Levels of solar radiation and other weather 
variables associated with dehydration on a given day 
were positively correlated with dew-drinking rate on 


VOLUME 56, NUMBER 2 


95 


TABLE 5. Three hypotheses regarding monarch butterfly early morning dew-drinking. 


Hypothesis Description 


Predictions 


(1) Current weather constraint 


(2) Previous day weather 


(3) Previous day mating activity 


among monarchs and should influence dew- 


drinking on the following morning. 


the following morning, but the effect was much more 
pronounced for males than females (Table 4b). 

A third hypothesis that may account for the dew- 
drinking pattern reported here relates to the level of 
mating activity on the previous day (Table 5—hypothe- 
sis 3). Similar to other butterflies, monarch mating 
consists of several stages (Rutowski 1991). Initially 
males locate potential partners either by perching on 
sun-exposed vegetation and waiting for females to pass 
nearby or by patrolling the air space near roosting 
trees (Falco 1999). Both of these locating tactics ex- 
pose males to direct sunlight, dry air, and high wind 
speeds. These conditions are more dehydrating than if 
they remained clustered. Early morning dew drinking 
by males and females may be a direct response to wa- 
ter deficit incurred primarily during mating effort on 
the previous day. 

Water uptake by male monarchs may influence their 
reproductive success indirectly. The majority of fe- 
males mate multiple times at California overwintering 
sites (Hill et al. 1976, Leong et al. 1995, Frey et al. 
1998, Frey 1999). A pattern of sperm precedence ex- 
ists so that sperm from the most recent mating fertil- 
izes the majority of eggs (Karen Oberhauser pers. 
com.) and sperm can survive for several weeks (Ober- 
hauser 1997). A male’s reproductive success is proba- 
bly a function of his partner’s subsequent inter-mating 
interval. Females that receive relatively large sper- 
matophores have longer inter-mating intervals (Ober- 
hauser 1989) and are more likely to exhibit effective 
resistance to male mating attempts (Frey 1999). Early 
morning dew drinking may enable males to generate 
accessory fluids necessary to produce a relatively large 
spermatophore during a subsequent mating and thus 
indirectly reduce chances that his sperm will be dis- 
placed. It is possible that dew drinking in this case is 


Flight from overnight cluster aggregations is 1. Meadow use should be greater on mornings 
constrained by low temperatures (Masters et al. ; 
1988), thus variation in morning temperature 
influences early morning meadow visits. 
Weather conditions associated with dehydration 1. Days with high levels of evapo-transpiration, wind 
(e.g., evapo-transpiration) during the previous day 
influence dew-drinking on the following morning. 


that warm more quickly. 


bo 


. Male and female pattern should be similar 


velocity, and solar radiation should be followed by 
mornings with high levels of dew-drinking and 
vice versa. 


bo 


. Male and female pattern of dew use should be similar 


High levels of mating activity increase dehydration 1. Measures of mating effort at the population level 


should correlate with dew-drinking rate on the 

following morning. 

Males should be more strongly affected than females. 

. Per capita dew-drinking rate should increase 
seasonally for males but not for females. 


wo bo 


also strategic behavior rather than a simple response to 
water deficit. All three predictions of the “previous day 
mating activity” hypothesis, i.e., Table 5—hypothesis 3, 
were supported by our findings (see Results—Factors 
affecting meadow use). 

Males that visited the meadow exhibited morpho- 
metric characters that were more similar to early 
morning mating-pool individuals than those captured 
later during the morning from clusters. Dew-drinking 
males had lower body weight (dry-mass), smaller ab- 
domens, and higher wing damage (Table 3). Male 
monarchs attempting to mate differed similarly from 
clustering males at Mexico and California overwinter- 
ing sites (Van Hook 1996, Frey et al. 1998, Oberhauser 
& Frey 1999). Both dew-drinking and mating males in 
our study had relatively high moisture content (Table 
3). Sunny, drier, and windier days were followed by in- 
creased male dew drinking the next morning (Table 4). 
In addition per capita dew drinking increased season- 
ally for males (but not for females) and their per capita 
mating effort also has been found to increase season- 
ally at this site (Falco 1999, Frey 1999). This is an ex- 
pected pattern if water “lost” by males, both from in- 
creased exposure to desiccation during mating 
attempts as well as during spermatophore transfer, was 
replenished by dew drinking. On the other hand, 
many of the meadow-visiting males attempted to cap- 
ture females within minutes of drinking dew. This is 
consistent with the view that early-morning dew drink- 
ing is a strategic male activity with the potential to in- 
crease short-term reproductive success in the ways de- 
scribed above. 

Male-biased sex ratios similar to those reported here 
for dew-drinking are described for puddling activity in 
other Lepidoptera (Adler 1982, Adler & Pearson 1982, 
Boggs & Jackson 1991, Sculley & Boggs 1996, Beck et 


96 


al. 1999; but see Scriber 1987). Puddling is often 
viewed as differential foraging, which either supplies 
ionic or nutrient resources needed by males (Arms et 
al. 1974) or is a means for males to acquire resources 
that they transfer to females during copulation (Scul- 
ley & Boggs 1996). Alternatively, or in addition, male 
biased puddling in Lepidoptera could provide mois- 
ture for reproductive activities, as well as, mainte- 
nance, as outlined above for monarchs. Monarchs also 
puddle at Mexico overwintering sites (pers. obs.) but 
puddling has not been reported for monarchs in west- 
ern North America. 

Nectaring from nearby flowering plants has been 
observed for monarchs during the mass mating phase 
at a Mexico overwintering site (Alonso-Mejia et al. 
1997). Flower-visiting monarchs in Mexico were 
smaller (dry mass), had significantly less fat reserves 
and had relatively greater moisture levels than cluster- 
captured ones (58% vs. 54% respectively for moisture 
as a proportion of wet body weight; our calculations of 
data from Table 2 in Alonso-Mejia et al. 1997). The 
pattern for moisture content of these Mexico mon- 
archs is similar to those from our dew-drinking study 
and indicates that nectaring can also provide monarchs 
with a source of water. Few nectar sources however 
occur near the Pismo overwintering habitat and mon- 
archs are seldom observed nectaring during January 
and February (pers. obs.). Likewise, they seldom drink 
from open water that usually occurs in shaded areas of 
the habitat, suggesting that monarchs probably obtain 
most of their water from dew or water droplets. 

Dew drinking by monarchs, like nectaring and pud- 
dling, may be “triggered” by a number of proximate- 
level factors and may provide water that is used in sev- 
eral functional contexts. Early morning dew drinking 
by monarchs seems to be a response to short-term de- 
hydration as well as strategic behavior associated with 
male reproductive activity. 


ACKNOWLEDGEMENTS 


This work was supported by a California State Faculty Support 
grant to Dennis Frey. We are grateful to Califormmia Parks and Recre- 
ation Department for allowing us to conduct monarch research on 
their property. Monarchs were collected under a permit issued to 
Dennis Frey by the California Parks and Recreation Department. 
Special thanks go to Lincoln Brower and Lisa Falco who provided 
insight and suggestions regarding both the project and manuscript. 
We are also thankful for valuable reviews by Karen Oberhauser, 
Carla Penz, and an anonymous reviewer. 


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Received for publication 11 September 2000; revised and accepted 5 
December 2001. 


Journal of the Lepidopterists’ Society 
56(2), 2002, 98-103 


SIBLING RIVALRY IN FLORIDA: THE DISPLACEMENT OF PYRGUS COMMUNIS 
BY PYRGUS ALBESCENS (HESPERIDAE) 


JOHN V. CALHOUN! 
977 Wicks Drive, Palm Harbor, Florida 34684-4656, USA 


ABSTRACT. A total of 204 specimens were collected during a field survey of Pyrgus conducted in Florida during 1989-2001. Identifica- 
tion of males was based on genitalic examination. Males of Pyrgus albescens were recorded from 32 Florida counties. Males of Pyrgus commu- 
nis were recorded from only 6 counties. Based on the results of this survey, as well as a review of 86 male specimens in public and private col- 
lections dating 1895-2001, it can be concluded that only P. communis originally occurred in Florida, but has recently been displaced by P. 
albescens. Pyrgus albescens was also found in Alabama and Georgia, and may be approaching South Carolina. Presented are details of the field 
survey, as well as a listing of all specimens used in this study. Also provided is information on habitats and hostplants of P. communis and P. 


albescens in Florida. 


Additional key words: Alabama, distribution, drought, Georgia, habitats, hostplants. 


Burns (2000) clarified the status of the sibling 
species Pyrgus communis (Grote) and Pyrgus albescens 
Pl6tz (common checkered skipper and white check- 
ered skipper, respectively). There is no known reliable 
method to separate these species based on wing pat- 
tern, but male genitalia exhibit consistent differences 
in the shape of the distal end of the left valve (Burns 
2000). As a result of this study, P. albescens was shown 
to be much more widespread than previously believed. 
This species was once thought to be limited to the 
southwestern United States and Mexico, but is now 
known to range eastward across the Gulf Coast states 
to Florida. The Florida distribution of P. albescens re- 
vealed by Burns (2000) is largely the result of a contin- 
uing survey of Pyrgus I have conducted since 1989. 

In September 1989, I captured what appeared to be 
two male P. communis in Calhoun County of the 
Florida panhandle. Astonished by the scarcity of re- 
cent reports of P. communis in Florida, I decided to 
obtain voucher specimens whenever the species was 
encountered. In October of that year, I located a siz- 
able population of P. communis in Pasco County of 
central Florida. Upon learning that John M. Burns 
(pers. com.) had found P. albescens in the extreme 
western Florida panhandle five years earlier, I decided 
to examine my specimens more closely. While the 
Pasco County individuals were clearly P. communis, 
the genitalia of those from Calhoun County were sur- 
prisingly consistent with P. albescens. Further in- 
trigued, I continued to scrutinize the genitalia of all “P 
communis” I obtained. 

Nine additional male P. albescens were captured in 
late 1994 and early 1995 in Calhoun, Columbia, 
Hamilton, Jackson, and Liberty counties of northern 
Florida. Most of these specimens were forwarded to J. 
M. Burns who confirmed their identity. These records 
suggested that P. albescens was even more widely dis- 


‘Research Associate, Florida State Collection of Arthropods, DPI, 


FDACS, Gainesville, Florida 32614, USA. 


tributed in Florida. Since that time, I have continued 
to sample Pyrgus at every opportunity. As a result, I 
have found that P. albescens is expanding in Florida 
and has displaced P, communis in the process. Pre- 
sented here are details of this survey, a review of his- 
torical specimens, and information on the habitat and 
hostplants of P. albescens in Florida. 


MATERIALS AND METHODS 


Field surveys for P. communis and P. albescens were 
conducted in Florida during 1989-1992, 1994-1997, 
1999-2001. They included trips expressly to locate 
Pyrgus, as well as opportunistic sampling during other 
research projects. Suitable habitats were identified via 
automobile and investigated on foot. Site visits were 
typically one hour or less in duration, depending upon 
site size (some were little more than narrow roadsides, 
others were multi-hectare pastures) and abundance of 
adults (fewer adults required more search time). If 
adults were not found, searches were discontinued af- 
ter 30 minutes. When populations were located, males 
were randomly collected and the genitalia examined 
by brushing away the scales from the left valves under 
a stereomicroscope. Females were also obtained, but 
they cannot reliably be separated (Burns 2000). Most 
females were tentatively determined by association 
with identified males. The remaining females were not 
assigned to either species. Because females are insep- 
arable, none were considered when evaluating the dis- 
tributions of P. albescens and P. communis in Florida. 

Also examined were Florida specimens deposited in 
various public and private collections. Male were de- 
termined through genitalic examination as follows: 
specimens in the National Museum of Natural History, 
American Museum of Natural History, and Florida 
State Collection of Arthropods were identified by J. M. 
Burns, specimens in The Natural History Museum 
(London) were identified by Kim Goodger; the single 
1978 specimen in the Allyn Museum of Entomology 
was identified by J. Y. Miller; T. M. Neal and A. D. 


VOLUME 56, NUMBER 2 


Fic. 1. Locations of male Pyrgus captures during the period 
1895-2001, where circles represent the 1989-2001 field survey and 
triangles are specimens in public and private collections. Solid circles 
and triangles, P. albescens; open circles and triangles, P. communis; 
half-solid circles and triangle, both species. Solid circles outside 
Florida represent survey specimens of P. albescens from Houston 
Co., Alabama, and Seminole Co., Georgia (USNM). 


Warren identified specimens in their personal collec- 
tions; all other specimens were identified by J. V. Cal- 
houn (JVC). 


RESULTS 


The field survey yielded a total of 204 specimens, 
obtained from 45 locations in 33 Florida counties (Fig. 
1, Table 1). Males of P albescens (n = 138) were 
recorded at 36 locations in 32 counties. Extreme col- 
lection dates range from § April-16 November. Males 
of P. albescens were also captured on 15 October 1995 
near Grangeburg in Houston Co., Alabama (n = 1) and 
near Riverturn in Seminole Co., Georgia (n = 3). In 
contrast, males of P communis (n = 15) were only ob- 
tained from 6 locations in 6 Florida counties. Extreme 
collection dates range from 16 July—31 December. Fe- 
males tentatively assigned to P. albescens (n = 31) were 
recorded at 12 locations in 12 counties. Females tenta- 
tively assigned to P. communis (n = 8) were recorded at 
3 locations in 3 counties. Unassigned females (n = 12) 
were recorded at 8 locations in 7 counties. One hun- 
dred and thirteen male and 11 female specimens col- 
lected between 1989 and 1999 were provided to J. M. 
Burns at the National Museum of Natural History 
(USNM), Washington, D.C. Most of the remaining 
specimens are deposited in my personal collection. 

Due to the general apathy exhibited by lepidopter- 


ists toward anything resembling the “common” P. com- 


99 


munis, relatively few Florida specimens exist in public 
and private collections. Nonetheless, 86 male speci- 
mens from Florida were ultimately located (Table 2). 
The majority of these specimens represent P. commu- 
nis (n = 56), collected in 15 counties between 1895 
and 1998. The remaining male specimens are P. 
albescens (n = 30), more recently collected in 7 coun- 
ties during the period 1976-2001. 

At no time have both P. albescens and P. communis 
been encountered together at the same location in 
Florida. However, P. oileus (Linnaeus) shares many lo- 
cations (and hostplants) with its congeners. Habitats 
for these species in Florida include vacant lots, weedy 
pastures, fallow cropland, farmyards, edges of culti- 
vated fields, open roadsides and citrus groves. These 
habitats are generally characterized by low-growing 
vegetation and an abundance of nectar sources, inter- 
spersed with patches of bare ground. Favorite flowers 
of both species are mostly white and include Bidens 
alba (L.) DC (Asteraceae), Phyla nodiflora (L.) 
Greene (Verbenaceae), Melilotus albus Medik. 
(Fabaceae), Richardia brasiliensis Gomez (Rubiaceae) 
and Sida spp. (Malvaceae). In Jackson County, 
Florida, I observed P. albescens ovipositing on Sida 
rhombifolia L., which also serves as a host of P. 
albescens and P. communis in Texas (Kendall 1965, 
Neck 1996, Burns 2000), as well as P. oileus in Texas 
and Florida (Kendall 1976, Minno & Emmel 1993). In 
1997 and 1999, Marc C. Minno reared P. albescens 
from larvae found on S. rhombifolia in Okaloosa and 
Brevard counties of Florida (adults det. by JVC). In 
Brevard County, I also found P. albescens in associa- 
tion with another, unidentified Sida species. It should 
be noted that some (or all) of the eight larval and two 
pupal specimens from Alachua County that Minno 
(1994) attributed to P communis could actually repre- 
sent P. albescens (no differences in the early stages of 
these species have yet been documented). 

Although Smith et. al (1994) observed that P. oileus 
and P. communis are indistinguishable on the wing, 
males of P. oileus appear whiter in color and both sexes 
of this species have a more sluggish, bobbing flight. 
Pyrgus oileus also tends to frequent semi-shaded situa- 
tions, whereas P. communis and P. albescens rarely 
stray from direct sunlight. Flight behavior of P. com- 
munis and P. albescens does not appear to differ. Both 
species fly rapidly near the ground and pause often to 
visit flowers. Males spend much time flying low cir- 
cuitous routes in search of females and will investigate 
virtually any movement, including other male Pyrgus, 
grasshoppers, and even falling leaves. This pugnacious 
behavior can become frustrating to anyone attempting 
to approach resting males, especially if grasshoppers 


100 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


TABLE 1. Pyrgus albescens and P. communis records documented in Florida during 1989-2001 field survey. AME, Allyn Museum of Ento- 
mology, JVC, John V. Calhoun; USNM, National Museum of Natural History. 


Nearest Speci- Nearest Speci- 
Date County town/city mens Collection Date County town/city mens Collection 
1. Pyrgus albescens (males) 1.ix.90 Pasco Dade City 2 USNM 
4 viii.94 Columbia Lake City 2 USNM 8.ix.90 Polk Branchborough 2 USNM 
4 viii.94 Hamilton Jasper 1 Jvc 30.ix.90 Pinellas Tarpon Springs 2 USNM/JVC 
8.iv.95 Calhoun Altha 2 USNM 23.xi.90 Pasco Dade City 2 USNM/AME 
8.iv.95 Jackson Oakdale 3. USNM 31.xii.90 Pasco Dade City 2 USNM 
8.iv.95 Liberty Bristol 1 USNM 16.vii.94 Hernando Brooksville 1 USNM 
15.x.95 Calhoun Altha 5 USNM 27 .viii.95 Lake Clermont 1 USNM 
15.x.95 Jackson Malone 1 USNM Total 15 
15.x.95 Jackson Oakdale 5 USNM 
30.x.95 Hamilton Jasper 4 (lost in post) 3. P. albescens (assigned females) 
30.x.95 Suwannee Pouchers 1 (lost in post) 4 viii.94 Columbia Lake City 2. JVG 
Corner 15.x.95 Jackson Oakdale 6 USNM 
5.ix.96 Jackson Marianna i MC 12.x.96 Jackson Oakdale 3) Ive 
12.x.96 Gadsden Rosedale 1 USNM 13.x.96 Jackson Oakdale 2 jvc 
12.x.96 Jackson Oakdale 4 JVC 3.ix.99 Alachua Alachua 1 Jvc 
13.x.96 Jackson Marianna 2 JVC 23.ix.99 Brevard Scottsmoor 2 JVC 
29.vi.97 Jackson Greenwood 1 jvc 23.ix.99 Volusia Scottsmoor 2 JVC 
3.ix.99 Alachua Alachua 2  USNM/JVC 1.x.99 Jefferson Lamont 2 Jvc 
23.ix.99 Brevard Scottsmoor 6 USNM 2.x.99 Santa Rosa Harold 3 ve 
23.ix.99 Volusia Scottsmoor 5 USNM 8.xi.00 Hillsborough Piney Point 2 Jvc 
1.x.99 Gilchrist Wilcox 7 USNM 16.ix.01 Lee Alva 2 ve 
1.x.99 Levy Chiefland 5 USNM 7.xi.01 Hemando Rital 1 jve 
1.x.99 Jefferson Lamont 5 USNM 7.xi.01 Polk Kathleen Ive 
1.x.99 Holmes Ponce de Leon 4  USNM 16.xi.01 Marion Marion Oaks 2 Jvc 
2.x.99 Okaloosa Cotton Bridge 7 USNM Total : 
2.x.99 Santa Rosa Harold 4  USNM 
2.x.99 Walton Mossy Head 3 USNM 4. P. communis (assigned females) 
3.x.99 Bay Saunders 5 USNM 29.x.89 Pasco Dade City Jvc 
3.x.99 Franklin Apalachicola 2 USNM L.ix.90 Pasco Dade City Jvc 
3.x.99 Gulf Wewahitchka 3. USNM 8.ix.90 Polk Branchborough Jvc 
3.x.99 Washington Orange Hill 1 USNM 30.ix.90 Pinellas Tarpon Springs JVC 
Corners 1.x.90 Pinellas Tarpon Springs JVC 
11.x.99 Pasco Dade City 2 USNM 23.xi.90 Pasco Dade City Jvc 
8.ix.00 Hillsborough Gulf City 1 Jve 29.ix.91 Pinellas Tarpon Springs Jvc 
1L.v.01 Lafayette Grady 1 jvc Total 
16.ix.01 Manatee Piney Point 3. JVC 
16.ix.01 Lee Alva 2 JVC 5. Pyrgus (unassigned females) 
20.ix.01 Pinellas Tarpon Springs 7 JVC 23.ix.90 Citrus Chassahowitzka 1 USNM 
7.xi.01 Citrus Bay Hill 1 jVve 27.viii.91 Pasco Dade City 2 JVC 
7.xi.01 Hernando Rital 3. JVC 30.viii.92 Pasco Dade City 1 jvc 
7.xi.01 Polk Kathleen 1 JVC 23.ix.92 Levy Yankeetown 1 JVC 
7.xi.01 Sumter Nobleton 2 JVC 8.ix.94 Pasco Dade City 1 Jvc 
16.xi.01 Marion Marion Oaks 2 Jvc 16.x.94 Volusia Scottsborough 2  USNM/JVC 
Total 138 13.xi.94 Hermando Dixie 1 USNM 
14.x.95 Jefferson Lamont 2 USNM 
2. P. communis (males) 12.x.96 Gadsden Chatahootchee 1 Jvc 
30.ix.89 Gadsden Concord USNM Total 12 
29.x.89 Pasco Dade City USNM 


are flushed with every footstep. Males also perch on 
taller vegetation, permitting them to observe and ex- 
amine passing objects easily. Adults of P. albescens and 
P. oileus have been seen roosting for the night on ex- 
posed herbaceous growth with wings tightly closed. 
This posture probably provides maximum solar expo- 
sure the following morning. Pyrgus albescens and P. 


communis reach maximum abundance during Septem- 
ber-November, when Sida hosts are plentiful. Al- 
though P. albescens and P. communis can be locally 
common where found, abundance can vary consid- 
erably between sites. Few adults were observed at 
many locations, accounting for the numerous single- 
specimen records documented during my survey. 


VOLUME 56, NUMBER 2. 


DISCUSSION 


Pyrgus communis has been reported from 41 
Florida counties, but many records are based on ob- 
servations and literature where specimens are unavail- 
able or lost (unpublished obs.). A number of literature 
reports are referable to P. oileus, especially females. 
For example, Grossbeck (1917) and Kimball (1965) 
listed P. communis specimens of W. T. Davis from Key 
West (Monroe Co.), Lakeland (Polk Co.) and Jack- 
sonville (Duval Co.) that are now deposited in the 
Staten Island Institute of Arts and Sciences (det. by 
JVC). The two Key West specimens (16.ix.1913) are 
female P. oileus, confirming the suspicions of Minno 
and Emmel (1993). The two specimens from Lakeland 
(8.xi.1913) and Jacksonville (7.xi.1913) are likely fe- 
male P. communis. Brewer (1982) listed P communis, 
but not P. oileus, from Sanibel Island, Lee County. 
However, her local collection deposited at the Sanibel- 
Captiva Conservation Foundation contains just the op- 
posite (P. oileus, but no P. communis). 

Pyrgus albescens may have been present in Florida 
for some time, sustaining small, highly localized (i.e., 
easily overlooked) populations that suddenly expanded 
due to unknown reasons. Alternatively, the species 
spread eastward around the Gulf of Mexico into 
Florida where it rapidly dispersed across the panhan- 
dle, then southward through the peninsula. Historical 
specimens further support the more likely scenario 
that P. albescens has only recently invaded the state. 

Based on specimens obtained during my field sur- 
vey (Table 1), as well as those from other collections 
(Table 2), it can be concluded that only P. communis 
originally occurred in Florida. The first known male P. 
albescens specimen from Florida was collected in 1976 
in Escambia County in the extreme western panhan- 
dle. All 54 male specimens collected during the 90 
years prior to 1976 are P. communis. In 1984, J. M. 
Burns found additional P. albescens at another location 
in Escambia County. By 1992, this species had reached 
Gadsden County in the eastern panhandle. The last P. 
communis collected in the panhandle was in 1989. All 
72 males collected after 1989 at 20 locations in 13 
counties throughout the panhandle are P. albescens, 
thus this species has probably dominated that region 
since at least the late 1980's or early 1990's. By 1994, P 
albescens had reached eastward in northern Florida to 
Columbia County and southward in the peninsula to 
Lake County. In 2001, P. albescens was found as far 
south as Lee and Okeechobee counties. The last con- 
firmed P. communis recorded in Florida was in 1998 in 
Levy County of the northwestern peninsula. Since that 
time, all 72 males collected at 21 locations in 17 coun- 


101 


ties of the peninsula (including Levy Co.) represent P 
albescens. The paucity of P. communis populations 
found during my field survey suggests that the ex- 
pansion of P. albescens in Florida had begun prior to 
1989. 

The southward progression of P. albescens through 
peninsular Florida, and associated displacement of P. 
communis, is reflected by several records. In 1994 and 
1995, I collected single males of P. communis (no P. 
albescens) in Hernando and Lake counties of the cen- 
tral peninsula. Also in 1994, D. R. Fine captured a 
single male P. albescens (no P. communis) at a more 
northern location in Lake County, suggesting this 
species was just invading that region. Evidence of di- 
rect displacement of P. communis by P. albescens was 
documented at three locations in northern and central 
Florida. In 1999, I captured only P. albescens in an 
agricultural field in Pasco County where only P. com- 
munis was recorded in 1989-1990. Likewise, in 2001, 
I found only P. albescens in a Pinellas County pasture 
where only P. communis had been collected in 1990. 
Unfortunately, dates of capture at these locations are 
nine or ten years apart, making the actual time of dis- 
placement difficult to determine. However, additional 
records from Levy County ostensibly limit displace- 
ment at one location to within six months. 

On 4 October 1998, Ron Hirzel collected 2 male P 
communis, but no P. albescens, at the crossroads town 
of Gulf Hammock in southern Levy County (Table 2). 
On 15 April 1999, and 20 March 2000, D. R. Fine cap- 
tured 4 male P. albescens, but no P. communis, in the 
same area of Gulf Hammock. In 2001, Richard A. An- 
derson obtained another male P. albescens (no P. com- 
munis) at the same Gulf Hammock location. In Octo- 
ber 1999, I found only P. albescens at a site in Levy 
County approximately 27 km north of Gulf Hammock 
(Table 1). Although P. albescens already occurred 
much further southward at that time, small peripheral 
populations of P. communis like that at Gulf Hammock 
may not have been as quickly impacted. 

Although P. communis is considered rare in south- 
em Florida, P. albescens may prove more successful at 
colonizing this region. The only known specimens of P. 
communis from southern Florida are a single old male 
from Punta Gorda (Charlotte County, ca. 1930) and 
another male collected in Miami (Miami-Dade 
County) in 1946 (Table I). The late John L. Heinrich 
(in litt. 30 November 1988) reported PB. communis 
from Lee County, but only P. oileus are currently de- 
posited in his collection at the Calusa Nature Center 
and Planetarium in Fort Myers, Florida. In 2001, I 
found P. albescens, but no P. communis, in Lee County 
where I had encountered only P. oileus between 1976 


102 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


TABLE 2. Male P. albescens and P. communis specimens from Florida in public and private collections. AME, Allyn Museum of Entomology; 
AMNH, American Museum of Natural History; BMNH, The Natural History Museum (London); CMNH, Camegie Museum of Natural His- 
tory; DRF, David R. Fine; FSCA, Florida State Collection of Arthropods; JVC, John V. Calhoun; MCM, Marc C. Minno; RLB, Robert L. 


Beiriger, USNM, National Museum of Natural History. 


Speci- 
Date County Location mens Collection 
1. Pyrgus communis (males) 
v.&vi.1895 Seminole Sanford 1 BMNH 
19P? Alachua Gainesville 2 FSCA 
18.iv.19?? Charlotte Punta Gorda 1 USNM 
26.ix-2.x.14 Alachua Gainesville 1 AMNH 
4-8.x.14 Jefferson Monticello 1 AMNH 
?.xi 17 Marion Ocala 2 CMNH 
<1919 P “Florida” 2 BMNH 
<1939 P “Florida” 1 BMNH 
3.v.42 Alachua Gainesville 1 FSCA 
T.vi.43 Alachua Gainesville 1 FSCA 
11.vi.43 Alachua Gainesville 1 FSCA 
17.vi.43 Alachua Gainesville 1 FSCA 
6.vii.43 Alachua Gainesville 1 FSCA 
4 viii.43 Alachua Gainesville 1 FSCA 
25.iv.44 Alachua Gainesville 1 FSCA 
2.v.44 Alachua Gainesville 1 FSCA 
8.v.46 Miami-Dade Miami 1 AME 
28.11.49 Escambia Pensicola Nav. 1 USNM 
Air. Sta. 
24.ix.49 Escambia Perdido Bay 1 USNM 
4 vii.59 Duval Jacksonville 1 FSCA 
23.viii59 Duval Jacksonville 1 FSCA 
22,.xi.60 Duval Jacksonville 1 FSCA 
9.v.62 Clay Orange Park 1 FSCA 
3L.viii.63 Duval Jacksonville 2 FSCA 
19.x.63 Duval Jacksonville 5 FSCA 
14.ii.64 Duval Jacksonville 1 FSCA 
20.x.64. Duval Jacksonville 1 FSCA 
20.x.64 Clay Orange Park 1 FSCA 
1.1.68 Duval Jacksonville 1 FSCA 


and 1987 (Calhoun 1987). Smith et. al (1994) sought P 
communis in southern Florida without success, stating 
“the failure of this Nearctic butterfly to enter the ‘trop- 
ical’ zone of the peninsula is remarkable.” On 30 Au- 
gust 2001, 23 adults of the “common checkered skip- 
per’ were observed near Flamingo, at the very southern 
tip of Florida, within Everglades National Park (Miami- 
Dade County) (Linda & Buck Cooper pers. com.). On 
13 November 2001, another individual was observed 
in extreme southwestern Florida within Fakahatchee 
Strand State Preserve in Collier County (R. L. Emmitt 
pers. com.). Future research may confirm the suspi- 
cion that these populations represent P. albescens, thus 
confirming its complete penetration of Florida. 
Credible observations and photographs of “common 
checkered skippers” in Florida have become much 
more frequent within the past two years. It seems 
likely that these reports represent P. albescens and the 
species is successfully colonizing areas not previously 
occupied by P. communis. I personally encounter P. 


Speci- 
Date County Location mens Collection 
15.viii.68 Liberty Torreya State Park 1 FSCA 
18.ix.68 Liberty Sweetwater Creek 2 FSCA 
5.iv.69 Alachua Gainesville 2 FSCA 
6.iv.69 Alachua Gainesville 1 FSCA 
15.viii.73 Gadsden Quincy 1 TMN 
19.viii.73 Gadsden Quincy 3 TMN 
20.x.73 Duval Jacksonville 1 FSCA 
25 .vii.74 Alachua Gainesville 1 TMN 
31L.x.77 Duval Jacksonville 2 FSCA 
28 .iv.78 Franklin Apalachicola 1 AME 
13.vii.87 Lake Sugarloaf Mtn. 1 MCM 
9 11.96 Putnam Caravelle Ranch 
WMA il MCM 
4.x.98 Levy Gulf Hammock 2 ADW 
Total 56 

2. P. albescens (males) 
22.v.76 Escambia Pensacola Beach 1 USNM 
7.ix.84 Escambia Cantonment 12 USNM 
26.viii.92 Gadsden SW of Quincy 1 RLB 
P.vii.94 Lake Paisley 1 DRF 
5.x.97 Okaloosa Blackwater Riv. 

St. For. 1 MCM 
15.iv.99 Levy Gulf Hammock 2 DRF 
14.x.99 Brevard Moccasin Is. WMA 6 MCM 
20.ii1.00 Levy Gulf Hammock 2 DRF 
L.iv.01 Lake Paisley 1 DRF 
7.viii.01 Okeechobee — Hilolo 1 DRF 
2.ix.01 Levy Gulf Hammock 2 Jvc 

Total 30 


albescens in many habitats where only P. oileus was 
formerly observed. Pyrgus albescens also appears to be 
spreading northward. On 4 June 2000, D. R. Fine col- 
lected two males and one suspected female of this 
species (det. by JVC) at Darien along the Altamaha 
River in McIntosh County of eastern Georgia. This 
record, along with my 1995 specimens of P. albescens 
from Houston County, Alabama and Seminole County 
Georgia, may show this species is expanding through- 
out the southeast and may reach South Carolina in the 
near future (or has already done so). 

No other species of butterfly in Florida has so deftly 
displaced another. Recent invasions of Florida by Ur- 
banus dorantes Stoll (Hesperiidae) and Danaus eres- 
imus (Cramer) (Nymphalidae) are excellent examples 
of successful widespread colonization (Knudson 1974, 
Calhoun 1996). However, neither of these exotic 
species has noticeably impacted its resident congeners, 
Urbanus proteus (L.) (Hesperiidae) and Danaus gilip- 
pus (Cramer) (Nymphalidae). The reasons behind the 


VOLUME 56, NUMBER 2, 


incursion of P. albescens into Florida are baffling, but 
changes in precipitation levels may offer an enticing 
explanation. 

Tilden (1965) and subsequent authors (e.g., Mac- 
Neill 1975, Orsak 1978) associated P. albescens with 
hot, arid lowland climates. In Texas, Neck (1996) sim- 
ilarly associated P. communis with “cooler, moister 
northern habitats” and P. albescens with “warmer, drier, 
southern habitats.” Although Florida can scarcely be 
described as “arid,” much of the state experienced 
moderate to extreme drought during the last decade, 
especially in 1989-1990 and 1998-2000 (NCDC 
2001). These conditions, more pronounced in the 
peninsula, continued into 2001. Drought has been de- 
fined as “a condition of moisture deficit sufficient to 
have an adverse effect on vegetation, animals, and man 
over a sizeable area” (Warwick 1975). While Burns 
(2000) doubted the strict affinity of P. albescens to drier 
climates, the rapid expansion of this species in Florida 
during the 1990's may, at least in part, be a consequence 
of these drought conditions. Burns and Kendall (1969) 
suspected humidity to be a limiting factor in the distri- 
butions of closely related Pyrgus philetas Edwards and 
P. oileus in southwestern North America. 

Pyrgus communis and P. albescens do not appear to 
coexist at any location in Florida, but they do occur to- 
gether in many areas of the southwestern United 
States (Tilden 1965, Austin 1986, Burns 2000). The ex- 
pansion of each species in that region seems to be in- 
hibited by the other species (Burns 2000). However, 
competitive pressures and other limiting factors may 
differ in the southeastern United States where P. com- 
munis may not have historically interacted with P. 
albescens. The dynamics of these species within re- 
gions of sympatry are obscure and unpredictable. Over 
time, P. communis may rebound in Florida. Shifts in 
dominance between these species at a single site in 
Arizona occurred over periods as short as 26 days (J. 
Burns pers. com.). It is also plausible that P. albescens 
was once abundant in the southeast during the more 
distant past, but P. communis reasserted its dominance 
until recently. Continued monitoring of Pyrgus may 
reveal more about the intriguing relationship between 
these sibling species and the factors responsible for 
this extraordinary displacement. 


ACKNOWLEDGMENTS 


Thanks are extended to John M. Burns (National Museum of 
Natural History) for many enjoyable and motivating conversations 
about Pyrgus and Hesperiidae in general. John M. Burns, Jim Cox 
(Tall Timbers Research Station), John B. Heppner (Florida State 
Collection of Arthropods), Kim Goodger (The Natural History Mu- 
seum, London), Ed Johnson (Staten Island Institute of Arts and Sci- 
ences), Jacqueline Y. Miller and Lee D. Miller (Allyn Museum of 


103 


Entomology), Philip D. Perkins (Museum of Comparative Zoology), 
Eric L. Quinter (American Museum of Natural History), John E. 
Rawlins (Carnegie Museum of Natural History), Dee Serage (Sani- 
bel-Captiva Conservation Foundation), and Laura Wewerka (Calusa 
Nature Center and Planetarium) provided information and access to 
specimens in their care. Richard A. Anderson, Robert L. Beiriger, 
David R. Fine, Mare C. Minno, Thomas M. Neal, Jeffrey R. Slotten, 
and Andrew D. Warren offered specimens and data from their per- 
sonal collections. Linda and Buck Cooper, Randy L. Emmitt, and 
Mark H. Salvato shared observations. John M. Burns and Andrew 
D. Warren critically reviewed the manuscript and provided helpful 
suggestions. 


LITERATURE CITED 


AusTIN, G. T. 1986. Pyrgus communis and P. albescens (Hesperi- 
idae) in Nevada. J. Lepid. Soc. 40:55-58. 

BREWER, J. 1982. A butterfly watcher’s guide to the butterflies of 
Sanibel and Captiva. Sanibel-Captiva [Florida] Cons. Founda- 
tion. 41 pp. 

Burns, J. M. 2000. Pyrgus communis and Pyrgus albescens (Hes- 
periidae: Pyginae) are separate transcontinental species with 
variable but diagnostic valves. J. Lepid. Soc. 54:52-71. 

Burns, J. M. & R. O. KENDALL. 1969. Ecologic and special distri- 
bution of Pyrgus oileus and Pyrgus philetas (Lepidoptera: Hes- 
periidae) at their northern distributional limits. Psyche 76:4-53. 

CALHOUN, J. V. 1988. A partial list of the butterflies and skippers of 
Lee County, Florida. S. Lepid. News 10:25-28. 

. 1996. Conquering soldiers: the successful invasion of 
Florida by Danaus eresimus (Lepidoptera: Nymphalidae). Hol- 
arctic Lepid. 3:7-18. 

GrOSSBECK, J. A. 1917. In F. E. Watson (ed.), Insects of Florida. IV. 
Bull. Amer. Mus. Nat. His. 37:1-147. 

KENDALL, R. O. 1965. Larval food plants and distribution notes 
from twenty-four Texas Hesperiidae. J. Lepid. Soc. 19:1-33. 

. 1976. Larval foodplants for thirty species of skippers (Lep- 

idoptera: Hesperiidae) from Mexico. Bull. Allyn Mus. No. 39. 9 


Mn C. P. 1965. Arthropods of Florida and neighboring land 
areas. Vol. I. Lepidoptera of Florida. Div. of Plant Ind. 363 pp. 

Knupbson, E. C. 1974. Urbanus dorantes dorantes Stoll (Hesperi- 
idae): another example of Florida’s population explosion. J. 
Lepid. Soc. 28:246-248. 

MacNEILL, C. D. 1975. Family Hesperiidae, pp. 423-578. In 
Howe, W. H. (ed.), The butterflies of North America. Double- 
day Page and Co., New York. 

MINNO, M. C. 1994. Immature stages of the skipper butterflies 
(Lepidoptera: Hesperiidae) of the United States: biology, mor- 
phology, and descriptions. Ph.D Dissertation, Univ. of Florida. 
509 pp. 

MINNO, M. C. & T. C. EMMEL. 1993. Butterflies of the Florida 
Keys. Scientific Publ., Gainesville, Florida. 168 pp. 

NCDC (NatIoNAL Ciimatic Data CENTER). 2001. US drought his- 
tory. National Oceanic and Atmospheric Admin. Web Page 
(Version 4 Oct 96). 
http:/Avww.ncdc.noaa.gov/climvis/usdroughthistory.html. 

Neck, R. W. 1996. A field guide to butterflies of Texas. Gulf Publ. 
Co., Houston, Texas. 323 pp. 

Orsak, L. J. 1978. The butterflies of Orange County, California. 
Univ. Calif. 349 pp. 

SmitH, D. S., L. D. MILLER & J. Y. MILLER. 1994. The butterflies 
of the West Indies and South Florida. Oxford Univ. Pr., New 
York. 264 pp., 32 pl. 

TILDEN, J. W. 1965. A note on Pyrgus communis and Pyrgus 
albescens (Hesperiidae). J. Lepid. Soc.19:91—94. 

WarWwICK, R. A. 1975. Drought hazard in the United States: a re- 
search assessment. Univ. Colorado, Inst. Bahavioral Sci. Mono- 
graph NSFRA/E-75/004. 199 pp. 


Received for publication 16 February 2001; revised and accepted 
24 November 2001. 


GENERAL NOTES 


Journal of the Lepidopterists’ Society 
56(2), 2002, 104-106 


IMMATURE STAGES OF SAIS ROSALIA (NYMPHALIDAE, ITHOMIINAE) 


Additional key words: life-history, Mechanitini, Solanaceae. 


The immature stages of butterflies in the subfamily 
Ithomiinae (Nymphalidae) are relatively well known 
for most genera, with good descriptions available in 
the literature (DeVries 1987, Brown & Freitas 1994 
and references therein). However, information is 
scarce or absent for many small genera (such as 
Roswellia, Eutresis, Athyrtis, Paititia, Aremfoxia, Vela- 
dyris, Velamysta, Dygoris, and Hypomenitis), and still 
incomplete for most larger diversified genera (Hya- 
lyris, Hypothyris, Napeogenes, Hyposcada, Oleria, 
Ithomia, Pteronymia, Greta, Hypoleria). In the tribe 
Mechanitini, there is adequate information for most 
species and all genera (Brown & Freitas 1994), except 
Forbestra (larva briefly described in Drummond 1976) 
and Sais (minimal information on the egg and larva in 
Table 1 and Fig. 2 of Brown & Freitas 1994, from rear- 
ing in Goiania, Goids, Brazil, in 1968). In this paper, all 
immature stages of Sais rosalia (Cramer, 1779) are de- 
scribed and illustrated. 

Sais rosalia (near subspecies rosalinde Weymer, 
1890 =paraensis Haensch, 1905, see Lamas 1994) was 
studied on banks and islands of the Teles Pires River, 
north of Alta Floresta, state of Mato Grosso, Brazil, in 
February and June 2000. On the latter visit, many eggs 
and larvae were collected on a solanaceous vine, prob- 
ably of the genus Lycianthes (very similar to the host 
recorded in Goiania; a live plant has been kept to await 
flowering). Additional eggs (n = 19) were also obtained 
from a wild caught female kept in a plastic bag with 
the host plant. The larvae were kept with leaves of the 
host plant in plastic boxes that were cleaned daily. Egg 
size was measured as height and width; and the larval 
head capsule size as the distance between the most ex- 
ternal ocelli (as in Freitas 1991, 1993); all capsules 
were retained for confirmation of growth stages. 
Adults, preserved larvae, capsules, and pupal skins are 
in the collection of the first author. 

About 20 eggs and four first instars were collected in 
the field on several food plant individuals growing near 
the riverbanks in sunny places. The plants varied from 
50 cm to 2 m high, and had small soft leaves. The iso- 
lated eggs and larvae were found on the underside of 
mature leaves. Females were observed ovipositing in 
the late afternoon, from 1500 to 1800 h. After inspect- 
ing the plant, the female landed on the upper side of a 
mature leaf and curved the abdomen to lay an isolated 
egg on the underside. After hatching, caterpillars ate 
part of the eggshell, and later began to eat the leaves, 


chewing small holes in the blade. Although solitary, 
larvae were not cannibalistic; several instars could be 
reared together without losses. The caterpillars rested 
in a J-shaped position on the underside of the leaves. 
When disturbed, caterpillars dropped off the leaf, sus- 
pending themselves by silk threads. 

Egg (Fig. la). White, elongated, slightly pointed at the apex, with 
15-18 longitudinal ridges and 9-12 transverse ridges (similar to that 
described in Brown & Freitas 1994). Duration 3-4 days. 

Larvae. First instar. White, turing green after first meal; legs, 
prolegs and anal plate black. Head black; average width 0.48 mm 
(SD = 0.03, n = 11). In dorsal view, the lateral tubercles (present in 
all instars) could be observed easily as small rounded projections on 
each abdominal segment. Maximum length 4 mm. Duration 2-3 
days. Second and third instars. Dark green to leaden gray, with a 
white collar on the prothorax and a lateral series of short yellow tu- 
bercles along the abdomen; legs, prolegs and anal plate pale. Head 
black; average width (second instar) 0.75 mm (SD = 0,03, n = 12), 
(third) 1.19 mm (SD = 0.04, n = 12). Maximum length 8 mm (sec- 
ond) or 13 mm (third instar). Duration 3 days (second) or 8-9 days 
(third). Fourth (final) instar (Fig. 1b, c). Dorsum dark gray, ventral 
region pale gray, with a white collar on the prothorax and a lateral se- 
ries of short yellow tubercles (tubercles on Al, A2, A7 and A8 more 
developed); legs and prolegs dark. Head black or rarely brown; aver- 
age width 1.74 mm (SD = 0.04, n = 10). Maximum length 25 mm. 
Duration 5-6 days. The pre-pupa loses the contrasting color-pattemm 
and becomes reddish, adopting an arched configuration (Fig. 1d). 

Pupa (Fig. le-g). Elongated, slightly arched ventrally (about 
60°) between the second and third abdominal segments, beige with 
a general gold reflection and black stripes on the wing cases (rarely 
dark with reduced reflective areas); cremaster red. Ocular caps short 
and pointed. Length 15 mm. Duration 13 days (n = 12). 

The immature stages of Sais rosalia are similar to 
those of other Mechanitini, including egg shape and 
size, number of longitudinal and transverse ridges, a 
lateral series of tubercles on the abdominal segments of 
the larvae, and the pupa elongated, slightly arched and 
reflective (Motta 1989, Brown & Freitas 1994). Con- 
trary to Mechanitis, which has long pointed lateral tu- 
bercles, the larva of Sais bears short lateral tubercles, a 
feature shared with Thyridia, Forbestra and Scada 
(Brown & Freitas 1994). Perhaps the most interesting 
feature in the life history of the studied population of 
Sais rosalia is that there were only four larval stadia. 
All known ithomiine larvae have at least five instars (six 
instars were observed once in Placidina, AVLF pers. 
obs.), with the exception of Tellervo zoilus (Fabricius, 
1775), also with four instars (Ackery 1987:272, with A. 
G. Orr); this Australian species has been placed in a 
separate subfamily by some authors (Ackery 1987). 

Only four instars were also observed in 12 larvae of 
Sais rosalia brasiliensis (reared to pupae on the Teles 


VOLUME 56, NUMBER 2 


105 


Fic. 1. Immature stages of Sais rosalia; a, egg; b, c, fourth (final) instar; d, pre-pupa; e-g, pupa (e, lateral; f, dorsal; g, ventral). 


Pires plant) from 17 eggs of a single female captured 
on 28 November 2001 in the broad floodable gallery 
forests of the Rio do Peixe (21°36’S, 51°42’W, 1400 
km south of the Teles Pires site, near Presidente 
Epitacio in extreme western Sao Paulo State). The re- 
covered head capsules from larvae reared in 2001 
show four widely separate width ranges: (1) 0.48-0.51 


mm (N = 7), (2) 0.74—0.77 mm (N = 5), (3) 1.15-1.24 
mm (N = 10, and (4) 1.72-1.96 mm (N = 8). All other 
aspects of larval coloration, instar duration, morphol- 
ogy and behavior were identical to the Teles Pires 
population. 


We are very grateful to the personnel of the Floresta Amaz6nica 
Hotel and the Rio Cristalino Jungle Lodge in Alta Floresta, MT 


106 


(Vitoria Da Riva Carvalho and her son Edson), for extensive and 
patient logistic support, housing and food during work in this re- 
gion in February and June 2000. Gerardo Lamas, Annette Aiello 
and Carla Penz made valuable comments on the manuscript. This 
research was partly supported by fellowships from the Brazilian 
Conselho Nacional de Desenvolvimento Cientifico e Tecnolégico 
(CNPq) and the Fundagaéo de Amparo a Pesquisa do Estado de 
Sao Paulo (BIOTA/FAPESP program, grants 98/05101-8 and 
00/01484-1). 


LITERATURE CITED 


ACKERY, P. R. 1987. The danaid genus Tellervo (Lepidoptera: 
Nymphalidae)—a cladistic approach. Zool. J. Linn. Soc. 
89:273-294. 

BROWN Jr., K. S. & A. V. L. Freitas. 1994. Juvenile stages of 
Ithomiinae: overview and systematics (Lepidoptera: Nymphali- 
dae). Trop. Lepid. 5:9-20. 

DEVRIES, P. J. 1987. The butterflies of Costa Rica and their natural 
history. Papilionidae, Pieridae, Nymphalidae. Princeton Univ. 
Press, Princeton, New Jersey. 327 pp- 

DRUMMOND III, B. A. 1976. Comparative ecology and mimetic re- 
lationships of Ithomiine butterflies in Eastern Ecuador. Ph.D. 


Journal of the Lepidopterists’ Society 
56(2), 2002, 106-108 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Thesis, University of Florida, xvi + 361 pp. 

Freitas, A. V, L. 1991. Variagao morfolégica, ciclo de vida e sis- 
tematica de Tegosa claudina (Eschscholtz) (Lepidoptera, 
Nymphalidae, Melitaeinae) no estado de Sao Paulo, Brasil. 
Revta. bras. Ent. 35:301—306. 

. 1993. Biology and population dynamics of Placidula eu- 
ryanassa, a relict ithomiine butterfly (Lepidoptera: Ithomiinae). 
J. Lepid. Soc. 47:87-105. 

Lamas, G. 1994. Los Danainae e Ithomiinae descritos por R. 
Haensch (Lepidoptera: Nymphalidae). Shilap 22:271-297. 
Morta, P. C. 1989. Andlise filogenética de Ithomiinae (Lep.: 
Nymphalidae) com base nos ovos: Relagao com plantas hos- 
pedeiras. MS Thesis. Universidade Estadual de Campinas. 

Campinas, SP, Brazil. 


ANDRE V, L. FREITAS AND KEITH S. BROWN JR., Museu de 
Historia Natural and Departamento de Zoologia, Instituto de Biolo- 
gia, Universidade Estadual de Campinas, C.P. 6109, Campinas, Sao 
Paulo, 13083-970, Brazil. 


Received for publication 20 August 2001; revised and accepted 26 
November 2001. 


MATINGS WITHOUT SPERMATOPHORE TRANSFER AND WITH TRANSFER OF TWO 
SPERMATOPHORES IN CALLOPHRYS XAMI (LYCAENIDAE) 


Additional key words: 


In Lepidoptera, males normally transfer one sper- 
matophore during copulation (Drummond 1984). How- 
ever, some studies indicate that in some matings no 
spermatophore is transferred (although this does not 
necessarily means that the female is not inseminated; 
Drummond 1984). There are several possible explana- 
tions for this fact: (a) exhaustion of substances necessary 
for building spermatophores as a result of frequent mat- 
ing (Drummond 1984); (b) male or female disabilities, 
such as deformations in the genitalia or in the reproduc- 
tive tract resulting from disease or defective develop- 
ment; or (c) mate choice (rejection) after initiation of 
copulation by females (i.e., females somehow inhibit or 
prevent the transfer of spermatophores by certain 
males; Eberhard 1996) or by males (i.e., males avoid to 
transfer spermatophores to certain females). Mate re- 
jection could be achieved by interrupting copulations 
before successful spermatophore transfer; in this case 
copulations are expected to be of short duration 
(Cordero 1993). 

On the other hand, it has been found that sometimes 
males transfer more than one spermatophore in one 
copulation (Drummond 1984). This type of mating may 
be a male adaptation to sperm competition if the trans- 
fer of multiple spermatophores decreases female recep- 


spermatophore production, copulation, sexual selection. 


tivity during a longer period (in several species there is 
a negative correlation between female receptivity and 
the degree of distention of the corpus bursa) (Drum- 
mond 1984) or if permits the transfer of more sperm 
(for example, if spermatophores can contain only a cer- 
tain maximum amount of sperm). However, the transfer 
of multiple spermatophores may be disadvantageous for 
many species, since the last spermatophore needs to be 
at least partially digested before re-mating because 
sperm migration to the spermatheca requires proper 
alignment of the spermatophore tube with the ductus 
seminalis and this alignment is more difficult in the 
presence of another spermatophore (Drummond 1984, 
Simmons & Siva-Jothy 1998). Thus, an alternative hy- 
pothesis is that the transfer of more than one sper- 
matophore in one copulation is result of a male disabil- 
ity. 

The multiplicity of explanations, and the theoretical 
relevance of many of them, indicates that to report mat- 
ings in which no spermatophore is transferred and in 
which multiple spermatophores are transferred, as well 
as its possible causes, is important. During the course of 
three laboratory experiments on spermatophore pro- 
duction by males of the lycaenid butterfly Callophrys 
xami (Reakirt) (Cordero 1998), in which I observed 199 


VOLUME 56, NUMBER 2, 


copulations, I recorded five copulations in which no 
spermatophore was transferred and four in which more 
than one spermatophore was deposited in the female 
corpus bursa. I report these observations in Table 1. 
My observations suggest two possible causes of fail- 
ure in spermatophore transfer. First, three of the four 
timed copulations lasted less (between ~7 and 15 min) 
than the average duration (+standard error) of first 
copulations of the day recorded in the field (32.3 + 4.9 
min; Cordero 1993) or in the laboratory (26.2 + 7.1 
min, 32.9 + 9.8 min and 35.9 + 10.2 min, considering 
each experiment separately; Cordero 1998), suggest- 
ing possible “interrupted copulations” (Cordero 1993). 
However, only in one case I observed the female be- 
havior associated with such copulations. I have dis- 
cussed the possibility that mate choice (by females or 
males) after mating began may be the cause of inter- 
rupted copulations (Cordero 1993). Second, female 
deformation of the ductus bursa and corpus bursa may 
have precluded the transfer of a spermatophore in 
case D (Table 1). Lifelong male disabilities may be dis- 
carded in all cases since the virgin male (case A) trans- 
ferred one spermatophore in a posterior copulation, 
and the four previously mated males (cases B—E) had 
transferred one spermatophore in a previous mating 
and two of them in subsequent matings (Table 1). Al- 
though the possibility of male exhaustion of substances 
necessary for building a spermatophore cannot be dis- 
carded in the case of the male that mated twice in a 
day (case C), we have observed several second matings 
of the day and in all cases one spermatophore was 
transferred. From my observations, I cannot suggest 
possible causes for the cases in which two sper- 
matophores were transferred during one copulation. 
It is unlikely that the “abnormal” matings reported 
in Table 1 are purely a product of laboratory condi- 
tions. Experimental matings were performed in cylin- 
drical mesh cloth cages (58 cm height and 26 cm di- 
ameter) hung outdoors in the natural habitat of C. 
xami, by individuals born in captivity. These conditions 
are artificial because territorial behavior (matings in 
the field are always performed by territorial males; 
Cordero & Sober6n 1990, Cordero et al. 2000) and the 
initial aerial components of courtship (Cordero 1993) 
were prevented by the experimental method. How- 
ever, in captivity all the non-aerial phases of courtship 
and copulation itself are, as far as I can tell, similar to 
those of occurring in the wild (Cordero 1993). Fur- 
thermore, the duration of matings in captivity and in 
the wild was very similar (see previous paragraph). Fi- 
nally, the 190 experimental “normal” matings in which 
one spermatophore was transferred and the nine “ab- 


107 


TABLE 1. Male mating status previous to relevant copulation and 
mating duration (MD) in which no spermatophore was transferred 
and in which two spermatophores were transferred by male Cal- 
lophrys xami. 


Male MD 


mating status (min) Comments 


Copulation without spermatophore transfer (n = 5) 

A. Virgin 15 We tried to re-mate this male the 
same day without success; the next 
day he mated (31 min) and trans- 
ferred two spermatophores (see case 
G); in a third mating he transferred 
one spermatophore 

Previous mating seven days before; 
male transferred one spermatophore 
in his first mating 

Previous mating the same day; male 
transferred one spermatophore in his 
first and in three subsequent matings 
During four min the female was moy- 
ing as in the “interrupted copulations” 
reported in Cordero (1993); this male 
mated the previous day during 20 min 
and transferred an spermatophore 
The female had deformed ductus 
bursa and corpus bursa; this male 
transferred an spermatophore in his 
first (two days before) and in a subse- 
quent mating 


B. Mated once 10 
C. Mated once > 


D. Mated once eA 


E. Mated once 38 


Copulation with transfer of two spermatophores (n = 4) 

F. Virgin (n = 3) ~18, ~25, ? These males transferred an sper- 
matophore in their two, two and four 
posterior matings, respectively 


G. Mated once 31 Male of case A 


normal” copulations occurred under apparently similar 
conditions (in fact, all “abnormal” copulations oc- 
curred in days in which “normal” matings were also 
observed). 

Although matings without spermatophore transfer 
and matings in which more than one spermatophore is 
transferred may be adaptive for males or females, they 
have negative effects on the fitness of at least one of 
the sexes (see above). The strength of the selective 
pressures exerted by the causes of these types of cop- 
ulations is correlated with the frequency of such copu- 
lations. The nine “abnormal” copulations reported in 
Table 1 correspond to 4.5% of all copulations observed 
during the three experiments (n = 199); and the eight 
males involved correspond to 10.5% of all experimen- 
tal males (n = 76). 


I thank the valuable comments made to the manuscript by Carla 
Penz and an anonymous reviewer. 


108 


LITERATURE CITED 


CoRDERO, C. 1993. The courtship behavior of Callophrys xami 
(Lycaenidae). J. Res. Lepid. 32:99-106. 

. 1998. Ecologia del Comportamiento Sexual de los Machos 
de la Mariposa Callophrys xami (Lycaenidae), con Algunas 
Consideraciones Acerca de la Evolucion del Semen de los In- 
sectos. Doctoral Thesis, UACPyP/CCH, UNAM, México. 

CoRDERO, C., R. MACIAS & G. JIMENEZ. 2000. The number of cop- 
ulations of territorial males of the butterfly Callophrys xami 
(Lycaenidae). J. Res. Lepid. 35:78-89. 

CorRDERO, C. & J. SOBERON. 1990. Non-resource based territorial- 
ity in males of the butterfly Xamia xami (Lepidoptera: Ly- 
caenidae). J. Insect Behav. 3:719-732. 

DRUMMOND III, B. A. 1984. Multiple mating and sperm competi- 
tion in the Lepidoptera, pp. 291-370. In Smith, R. L. (ed.), 
Sperm competition and the evolution of animal mating systems. 
Academic Press, New York. 


Journal of the Lepidopterists’ Society 
56(2), 2002, 108-111 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


EBERHARD W. G. 1996. Female control. Sexual selection by cryptic 
female choice. Princeton University Press, Princeton, U.S.A. 

SIMMONS, L. W. & M. T. Stva-JoTHy. 1998. Sperm competition in 
insects: mechanisms and the potential for selection, pp. 
341-434. In Birkhead, T. R. & A. P. Mogller (eds.), Sperm com- 
petition and sexual selection. Academic Press, London. 


CARLOS CORDERO!, CEAMISH, Universidad Autonoma del Es- 
tado de Morelos, Av. Universidad 1001, Col. Chamilpa, Cuernavaca 
62210, Morelos, México. 


Received for publication 7 January 2001; revised and accepted 28 
November 2001. 


‘Current address: Departamento de Ecologia Evolutiva, Instituto 
de Ecologia, UNAM, Apdo. Post. 70-275, Ciudad Universitaria, 
Coyoacan 04510, México, DF, México. 


PAPILIO DEMOLEUS (PAPILIONIDAE) IN BORNEO AND BALI 


Additional key words: Malaysia, Indonesia, Malay Archipelago, invasion, deforestation. 


Papilio demoleus L. is widely distributed in the trop- 
ical and subtropical regions of Asian continent, Aus- 
tralia, and the islands of Taiwan, Hainan, New Guinea, 
and Lesser Sunda Islands (Sumba, Flores and Alor), 
but it had been lacking in Sumatra, Java, Borneo, the 
Philippines, and the Moluccas until its recent invasion 
of these islands (Corbet & Pendlebury 1978, 1992). 
Sumatra received ssp. malayanus Wallace from the 
Malay Peninsula and the Philippines ssp. libanius 
Fruhstorfer from Taiwan in the 1960-70’s (Jumalon 
1968, Hiura 1973, Miyata 1973, Tsukada & Nishiyama 
1980). These two subspecies can be easily discrimi- 
nated based on the wing markings; Fruhstorfer (1908), 
in his description of ssp. libanius, stated that the Tai- 
wanese specimens appear darker than the nominate 
subspecies specimens from “Tonkin.” The dark appear- 
ance of ssp. libanius is mainly due to the fact that the 
yellow spots in spaces la and 1b of forewing upper sur- 
face are always narrow, whereas these spots are always 
very broad in ssp. malayanus, though as was not men- 
tioned in the Wallace’s (1865) description of the sub- 
species (Figs. 1-4). 

The Malay subspecies was confirmed to have estab- 
lished its population in Java, supposedly having in- 
vaded from Sumatra during late 1980's (Kato 1989, 
Matsumoto & Noerdjito 1996). The species has also 
been found from Borneo. Otsuka (1988) illustrated a 
male and a female specimen of P. demoleus from 
Keningau, Sabah, without mentioning that the records 
were new in Borneo. Although Otsuka (1988) did not 
identify the subspecies, the illustrated specimens ex- 
hibited typical characters of ssp. libanius. Ishii (1987, 


1991) identified P. d. libanius from Sandakan, Sabah, 
and he mentioned (Ishii 1991) that he saw only a few 
individuals of this species in 1981, while he found 
many individuals of the same species in 1983, and sug- 
gested that the Taiwanese subspecies may have in- 
vaded from the Philippines during the early 1980's. 


TABLE 1. Year of first record and presumed origin of the Papilio 
demoleus populations recently established in the Southeast Asian Is- 


lands. 


Year of Presumed 

Island first record origin Literature 

Luzon 1967* Taiwan Jumalon (1968) 

Cebu 1968? Taiwan Hiura (1973) 

Leyte 1968? Taiwan Hiura (1973) 

Palawan 1969 Taiwan Hiura (1973) 

Negros 1969? Taiwan Miyata (1973) 

Mindanao 1969 Taiwan Miyata (1973) 

Mindoro 1971 Taiwan Hiura (1973) 

Talaud Unknown** Taiwan Tsukada & 
Nishiyama (1980) 

Sangihe Unknown** Taiwan Tsukada & 
Nishiyama (1980) 

Sula Unknown** Taiwan Tsukada & 
Nishiyama (1980) 

Borneo 1983 Taiwan Ishii (1987) 

1996 Malay Peninsula This study 

Sumatra Unknown** Malay Peninsula Tsukada & 
Nishiyama (1980) 

Java 1988* Malay Peninsula Kato (1989) 

Bali 1991 Malay Peninsula This study 


* Except old sporadic records which are unrelated to the present 
population (see Jumalon 1968, Moonen 1991 for further details). 

** Tsukada & Nishiyama (1980) first stated occurrence of the 
species in Sumatra without indicating earliest collection data. 


VOLUME 56, NUMBER 2 109 


Fics. 1-4. Papilio demoleus collected in Bomeo and Bali: 1, a male with characters of ssp. malayanus (Bukit Soeharto, East Kalimantan, 
16 Noy. 1998); 2, a male with characters of ssp. libanius (Bukit Soeharto, East Kalimantan, 23 Noy. 1998); 3, a female with intermediate char- 
acters between ssp. malayanus and ssp. libanius (Tawau, Sabah, 11 June 1996); 4, male P d. malayanus (Nusa Dua, Bali, 17 February 1991). 


110 


MALAY 
PENINSULA 


— ® 
S 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


PHILIPPINES 


IWENCINE, 


CEBU 


MINDANAO 


FLORES ALOR@ “6 
yu 


»? BCE F949 


SUMBA* «© @ 


Fic. 5. A map of Southeast Asia, showing islands recently invaded by Papilio demoleus. Arrows indicate presumable direction of the invasion. 


Furthermore, P. d. libanius has been also recorded 
from Talaud, Sangihe and Sula Islands of the Moluccas 
(Tsukada & Nishiyama 1980, Corbet & Pendlebury 

1992), again suggesting invasion via the Philippines 
(Table 1, Fig. 5). 

Recently, I confirmed P. d. malayanus had invaded 
into Borneo (including Indonesian Kalimantan) and 
Bali as described below. 

P. demoleus in Borneo. I collected four males and 
one female at Luasong (near Tawau) and one female at 
Tawau in Sabah, Malaysia, in 1996 (the specimens are 
kept at Luasong Forestry Centre, Luasong, Sabah). 
Hiroshi Makihara of Forestry and Forest Beaches Re- 
search Institute (FFPRI), Japan, also collected 18 
males and three females at Bukit Soeharto (near 
Samarinda), East Kalimantan, Indonesia, in 1998 and 
showed them to me for examination (the specimens 
are temporarily kept by Makihara and to be kept else- 
where in Indonesia in the future). Ten of these 27 
specimens looked like ssp. libanius (1 d Luasong, 19 


March 1996; 1 2 Tawau, 11 June 1996; 1 6 Bukit Soe- 
harto, 6 October 1998; 4 d Bukit Soeharto, 23 Novem- 
ber 1998; 2 3 Bukit Soeharto, 8 December 1998; 1 d 
Bukit Soeharto, 30 November 1998), eight looked like 
ssp. malayanus (1 d Luasong, 19 March 1996; 1 d Lu- 
asong, 25 May 1996; 1 d Bukit Soeharto, 12 January 
1999: 2 d Bukit Soeharto, 6 October 1998; 2 2 Bukit 
Soeharrto, 2 November 1998; 1 6 Bukit Soeharto, 9 
November 1998), and nine exhibited various interme- 
diate characters between the two (1 ¢ Luasong, | June 
1996; 1 ° Luasong, 28 May 1996; 1 6 Bukit Soeharto, 6 
October 1998; 1 d, 1 2° Bukit Soeharto, 6 November 
1998; 1 d Bukit Soeharto, 9 November 1998; 1 6 Bukit 
Soeharto, 16 November 1998; 2 6 Bukit Soeharto, 23 
November 1998), indicating that the population in Bor- 
neo is now a mixture of the two subspecies (Figs. 1-3). 

P. demoleus in Bali. I found two males of P. de- 
moleus collected by Keizi Kiritani at Nusa Dua, Bali, 
Indonesia, on 17 February, 1991, in the collection of 
National Institute of Agro-Environmental Sciences 


VOLUME 56, NUMBER 2 


(NIAES), Japan. These specimens had typical charac- 
ters of ssp. malayanus (Fig. 4). To my knowledge, this 
is the first record of the species from Bali. Although 
there has been no more information of this species in 
Bali, it seems likely that the species is established 
there. 

Factors favoring spread of P. demoleus. Papilio 
demoleus frequents in and around villages and urban 
areas. It is originally a species of monsoon regions and 
prefers open habitats to thick shadowy rain forests as 
used to dominate in Sumatra, Java, Bali, Borneo and 
the islands of the Philippines. Hiura (1973) suggested 
that this habitat preference is the reason why this 
species had never occurred in the rain forest regions in 
Sunda Land and the Philippines before, while recent 
large scale deforestation in the Philippines prepared 
suitable habitats for the species. I share the same view 
with him and believe that the same logic applies to the 
recent establishment of the P. demoleus populations in 
Sumatra and Borneo. In densely human populated 
Java and Bali, forests have been exploited earlier and 
suitable habitats for P. demoleus have long been preva- 
lent. However, Java and Bali had been far derived 
from the nearest P. demoleus population, until effects 
of extensive logging and consequent human activities, 
e.g., fire, shifting cultivation, illegal settlements, cattle 
breeding, etc. in the forests (or ex-forests) became 
conspicuous in Sumatra in the 1960—70’s. After this 
stage, Sumatra played a role of stepping stone for P. d. 
malayanus to invade into Java, and then Java played a 
similar role between Sumatra and Bali. The occur- 
rence of P. demoleus in Bali in 1991 indicates that the 
invasion into Bali took place almost immediately after 
its establishment in Java. The Malay subspecies in Bor- 
neo could have invaded either from Sumatra or Java, 
or directly from Malay Peninsula (Fig. 5). 

Then, a question arises; why the Australian sub- 
species (i.e., ssp. sthenelus Macleay of Australia and 
Sumba, ssp. sthenelinus Rothschild of Flores and Alor 
and ssp. novoguineensis Rothschild of New Guinea) 
would not spread? Food plant availability could be the 
factor favoring spread of the Asian subspecies, not the 
Australian subspecies. The Asian subspecies feed on 
oranges, Citrus hystrix DC., C. aurantifolia Swingle, 
C. amblyocarpa Ochse, etc. (Rutaceae; Corbet & 
Pendlebury 1992, Matsumoto & Noerdjito 1996) 
which are very commonly planted either as a crop or 
an ornament in towns, villages and illegal settlements 
in the forest areas. The Australian subspecies feed on 
wild leguminous plants of the genus Psoralea L..: i.e., P. 
tenax Lindl., P. patens Lindl., P. cinerea Lindl, P. leu- 
cantha F. Muell. and P. pustulata F. Muell. in Australia 
(Common & Waterhouse 1981) and P. badocana 


111 


Benth. in Papua New Guinea (Parsons 1999) (no food 
plant record available for the Lesser Sunda popula- 
tions). Although Psoralea plants occur in the Sunda Is- 
lands (Bentham & Mueller 1967), there has been no 
evidence to indicate increase of these plants favored 
by recent environmental changes. The Citrus feeding 
Asian subspecies, on the other hand, are now expand- 
ing its range in the Malay Archipelago, being favored 
by increase of habitats with artificially planted Citrus 
hosts. 


I thank H. Makihara of FFPRI and NIAES for giving opportu- 
nity for me to examine their specimens. Scott E. Miller and Carla M. 
Penz reviewed the manuscript and made helpful comments and sug- 
gestions. 


LITERATURE CITED 


BENTHAM, G. & F. MUELLER. 1967. Flora Australiensis: a descrip- 
tion of the plants of the Australian Territory. A. Asher & Co., 
Amsterdam. 521 pp. 

Common, I. F. B. & D. F WaTERHOUSE. 1981. Butterflies of Aus- 
tralia. Angus & Robertson, Sydney. 682 pp. + 20 pls. 

Corset, A. S. & H. M. PENDLEBURY. 1978. The butterflies of the 
Malay Peninsula. 3rd ed. (rev. by J. N. Eliot). Malayan Nature 
Society, Kuala Lumpur. 578 pp. + 35 pls. 

. 1992. The butterflies of the Malay Peninsula. 4th ed. (rev. 
by J. N. Eliot). Malayan Nature Society, Kuala Lumpur. 595 pp. 
+ 69 pls. 

FRUHSTORFER, H. 1908. Lepidopterologisches Péle-Méle. Ent. 
Zeitschr. 22:140-141. 

Hiura, I. 1973. Butterflies flying across the sea. Soju Shobo, Tokyo. 
200 pp. 

IsHm, M. 1987. Diapause potential in tropical Papilionids (Lepi- 
doptera: Papilionidae). Appl. Ent. Zool. 22:114-15. 

. 1991. Tropics for butterflies, pp. 59-84. In Hidaka, T. & 
M. Ishii (eds.), Animals and plants in Borneo—studying their 
lives in tropical forests. Tokyo Kagaku Dojin, Tokyo. 348 pp: 

Jumaton, J. N. 1968. A comment on the new papilionid from the 
Philippines. Ty6 to Ga 19:105-109. 

Kato, S. 1989. Notes on Papilio demoleus Linnaeus collected in 
Java, Indonesia (Lepidoptera, Papilionidae). Ty6 to Ga 
40:189-191. 

Matsumoro, K. & A. W. Noerpjito. 1996. Establishment of Pa- 
pilio demoleus L. in Java. J. Lepid. Soc. 50:139-140. 

Miyata, A. 1973. Notes on Princeps demoleus libanius Fruhstorfer 
in the Philippines. Ty6 to Ga 24:37-41. 

MOONEN, J. J. M. 1991. Papilio demoleus L. in Java (Lep.; Papil- 
ionidae). Ty6 to Ga 42:93-94. 

Otsuka, K. 1988. Butterflies of Borneo. Vol. 1. Tobishima Corp., 
Tokyo. x + 61 pp. (in Japanese) + xix + 61 pp. (in English) + 80 

ls. 

Pe M. 1999. Butterflies of Papua New Guinea. Academic 
Press, San Diego. 736 pp. + 136 pls. 

TsuKADA, E. & Y. NISHIYAMA. 1980. Butterflies of South East Asian 
islands. I. Papilionidae. Plapac, Tokyo. 459 pp. 

WALLACE, A. R. 1865. On the phenomena of variation and geo- 
graphical distribution as illustrated by the Papilionidae al the 

Malayan region. Transactions of the Linnean Society 25:1-71 + 
pls. I-VIIL. 


Kazuma Matsumoto, Tama Forest Science Garden, Forestry and 
Forest Products Research Institute, Todori-ch6, Hachidji, Tokyo 193- 
0843, Japan. 


Received for publication 7 February 2001; revised and accepted 
3 December 2001. 


Journal of the Lepidopterists’ Society 
56(2), 2002, 112-114 


NOTES ON THE NEVADA BUCK MOTH, HEMILEUCA NEVADENSIS (SATURNIIDAE) IN 
SOUTHERN MANITOBA 


Additional key words: Tachinidae, Leschenaultia, bur oak, Hemileucinae, Apoecilus. 


Observations on the Nevada buck moth, Hemileuca 
nevadensis Stretch, in southern Manitoba are re- 
ported, along with information regarding new predator 
and parasitoid associations for this species, as well as a 
new host plant association. In Manitoba, H. nevadensis 
has been collected in widely separated localities like 
Aweme, Selkirk, Victoria Beach, and Winnipeg. Lim- 
ited collection records indicate that the flight period in 
southern Manitoba begins in late August or early Sep- 
tember. Adults are day flying, with mating and oviposit- 
ing occurring the same day (Tuskes et al. 1996). Eggs 
are laid in rings around twigs of willows (Salix spp.) 
and trembling aspen (Populus tremuloides Michaux). 
Eggs overwinter and hatch early in the spring, almost 
synchronously with leaf flush. During the first three in- 
stars larvae are black in coloration and are gregarious 
(Tuskes et al. 1996). In southern Manitoba, pupation 
occurs in mid-summer (late-June to mid-July). Adults 
eclose approximately two months later. Little else is 
known about the biology of this species in southern 
Manitoba. 

In early June 1998, several students and faculty 
from the University of Manitoba Department of Ento- 
mology traveled to the Manitoba Tall Grass Prairie 
Preserve located approximately 10 km east of the town 
of Roseau River, Manitoba (49.06°N, 96.7°W), (Fig. 
1). This locality is characterized as having an abun- 
dance of grasses, willows and poplars. Predominant 
grasses include big bluestem, Andropogon gerardii 
Vitman and little bluestem, Schizachyrium scoparium 
(Michx.) Nash (Gramineae). The junior author ob- 
served several hundred clusters of unidentified larvae 
feeding on the foliage of Salix spp. and P. tremuloides. 
Casual observation at this locality for at least the two 
years previous did not detect larval clusters. Several 
larval clusters were collected and returned to the lab- 
oratory for determination. Larvae were reared to ma- 
turity and identified by the senior author as H. 
nevadensis. The junior author also collected one larval 
cluster from bur oak, Quercus macrocarpa Michaux. 
This plant has not been previously reported as a food- 
plant for H. nevadensis. The closely related H. maia 
(Drury) has been reported to feed on various species 
of oak. However, H. maia is not known to occur in 
Manitoba. Tuskes et al. (1996) cited a record of H. 
nevadensis larvae feeding on coastal live oak, Quercus 
agrifolia Nee, in California. 


On 2 July 1998, the authors returned to the site of 
the original larval collections. At this time most larvae 
were in sixth-instar and could be collected by the hun- 
dreds from willow bushes in the area. Larvae were less 
commonly found feeding on foliage of P. tremuloides 
and Q. macrocarpa. Larvae that were feeding on Q. 
macrocarpa were stunted and appeared to be lagging 
a full instar behind larvae feeding on the other food- 
plants. One larva feeding on willow was observed un- 
der attack by a stink bug, Apoecilus bracteatus (Fitch) 
(Hemiptera: Pentatomidae). This predaceous stink 
bug is known to attack a diverse range of soft-bodied 
herbivorous insects (Evans & Root 1980). Kendall & 
Peigler (1981) reported that larvae of Hemileuca grotei 
Grote and Robinson were attacked by Apoecilus cyni- 
cus (Say) (cited as Apateticus cynicus) nymphs and 
adults. 

Mature H. nevadensis larvae were noted to be poly- 
morphic in coloration. The color of the integument 
ranged from yellow to almost completely black. Sey- 
eral hundred larvae were collected and returned to the 
laboratory in an effort to obtain parasitoids as well as 
living Hemileuca pupae. Pupae were maintained at 
25°C and 14:10 photoperiod. Adults began emerging 
in the laboratory during late August 1998. Approxi- 
mately one-half of the pupae did not produce adults 
and remained in diapause until the following fall. 
Tuskes et al. (1996) reported that a portion of 
Hemileuca populations may overwinter as pupae. 

On 7 September 1998, the senior author returned to 
the collection locality with five virgin females. These 
females began calling during mid-morning, while en- 
route from Winnipeg. When the locality was being ap- 
proached, males began following the vehicle. As soon 
as the vehicle stopped, hundreds of males were ob- 
served arriving from all directions in an effort to locate 
and mate with the caged females (Fig. 2). When the 
cage was opened, allowing wild males access, copula- 
tion occurred within ten seconds. Shortly thereafter, 
the remaining males dispersed. 

The rate of parasitism of H. nevadensis larvae was 
estimated to be approximately 20%. Several hundred 
tachinid puparia were obtained. Puparia were stored 
in a 5°C chamber for approximately three months and 
afterwards maintained at 20°C and 16:8 photoperiod 
until emergence of adult flies. Adults were identified as 
Leschenaultia fulvipes (Bigot) (Diptera: Tachinidae). 


VOLUME 56, NUMBER 2 


113 


Fics. 1, 2. Hemileuca nevadensis at the Manitoba Tall Grass Prairie Preserve. 1, Collection locality habitat, 10 km East of Roseau River. 2, 
Wild Hemileuca nevadensis males attracted to caged virgin females. Both photos taken 5 September 1998 by DCH. 


This species has not been recorded as attacking H. 
nevadensis, but has been recorded from H. maia, H. 
grotei, and H. lucina (Arnaud 1978, Kendall & Peigler 
1981, Peigler 1994). Members of this genus lay mi- 


crotype eggs on foliage. Eggs hatch shortly after inges- 
tion by the host (Brooks 1946). A figure of this para- 
sitoid ovipositing at Hemileuca larvae is in Tuskes et al. 
(1996:26). Preserved egg rings, larvae and adults of H. 


114 


nevadensis, and adult Leschenaultia fulvipes were de- 
posited in the J. B. Wallis Museum of Entomology, 
University of Manitoba Department of Entomology. 
Identification of adult Hemileuca followed the de- 
scriptions of Ferguson (1971) and Tuskes et al. (1996). It 
should be noted here that Ferguson (1971) referred to 
the Manitoba populations of H. nevadensis as latifascia 
Barnes and McDunnouogh. 
Hemileuca near Roseau River, Manitoba seems to be 
somewhat enigmatic. The association of several larval 
clusters with Q. macrocarpa would appear to be a H. 
maia trait. Ferguson (1971) was unsure that H. nevaden- 
sis warranted treatment as a ‘full’ species, since he could 
not distinguish the genitalia from those of H. maia. Fer- 
guson (1971) also discussed the existence of transitional 
populations between maia and nevadensis in the Mid- 
west. The H. nevadensis population near Roseau River, 
Manitoba may represent one such transitional popula- 
tion. Tuskes et al. (1996) discuss the taxonomic problems 
associated with this genus in the Great Lakes region and 
assert that more research on Hemileuca populations in 
this region is needed to resolve their taxonomic status. 


The authors wish to thank J. E. O'Hara of Agriculture and Agri- 
Food Canada for identification of tachinid adults. Thanks are also 
extended to Dr. Abner Hammond, Louisiana State University, for 
helpful comments and criticisms of this manuscript. 


The population of 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


LITERATURE CITED 


ARNAUD Jr., P. H. 1978. A host-parasite catalog of North American 
Tachinidae (Diptera). United States Department of Agricul- 
ture, Miscellaneous Publication 1319:1—-860 + 1 pl. 

Brooks, A. R. 1946. A revision of the North American species of 
Leschenaultia Sens. Lat. (Diptera: Larvaevoridae). Can. Ent. 
78:169-182. 

Evans, E. W. & R. B. Roor. 1980. Group molting and other life 
ways of a solitary hunter, Apateticus bracteatus (Hemiptera: 
Pentatomidae). Ann. Entomol. Soc. Am. 73:270-274. 

FERGUSON, D. C. 1971. Fasc.20.2A, Bombycoidea (in part), pp. 
101-153. In R. B. Dominick et al., The moths of America north 
of Mexico, Fasc. 20.2A, Bombycoidea (in Part). Classey, Lon- 
don. 

KENDALL, R. O. & R. S. PEIGLER. 1981. Hemileuca grotei (Saturni- 
idae): its morphology, natural history, spatial and temporal dis- 
tribution. J. Lepid. Soc. 35:41—50. 

PEIGLER, R.S. 1994 (1996). Catalog of Parasitoids of Saturniidae of 
the World. J. Res. Lepid. 33:1-121. 

TuskEs, P. M., J. P. TurrLe & M. M. Couns. 1996. The Wild Silk 
Moths of North America. A natural history of the Saturniidae of 
the United States and Canada. Cornell University Press, Ithaca, 
New York. 250 pp. 


D. C. HENNE! AND J. K. DIEHL. Department of Entomology, 214 
Animal Science/Entomology, University of Manitoba, Winnipeg, 
Manitoba R3T 2N2, CANADA. 


Received for publication 8 January 2001; revised and accepted 10 
December 2001. 


' Present address: Department of Entomology, 402 Life Sciences 
Building, Louisiana State University Agricultural Center, Baton 
Rouge, iLomistena 70803, USA. 


Journal of the Lepidopterists’ Society 
56(2), 2002, 115 


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CATERPILLAR CALLS 


The illustration depicts a Eurybia patrona (Riodinidae, Eurybiini) caterpillar delivering food secretions to an ant 
from paired tentacle nectary organs located on the caterpillar’s eight abdominal segment. Drawing by J. Clark 
In: DeVries, P.J. 1997. Butterflies of Costa Rica and their natural history: Riodinidae. Princeton University Press. 

In addition to amino acid-rich secretions that reward symbionts in exchange for protection, myrmecophilous 
lycaenids and riodinids produce substrate-borne calls that attract and maintain ant guards. Call mechanisms vary 
within the Riodinidae, and between riodinids and lycaenids, as does the location and structure of the secretion-pro- 
ducing organs. Nonetheless, call production has been demonstrated as being of critical importance to the mainte- 
nance of a guard of ants. Although most caterpillars eat plant tissue (leaves or flowers), some myrmecophilous 
riodinids are free-living carnivores, and some lycaenids spend part of their life inside ant nests feeding on ant re- 
gurgitations and ant brood. 


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CONTENTS 


Winstow J. Howarp: PIONEER NEw Mexico naturauist Steven J. Cary ---------------------- === AQ Fiat 
FIVE NEW SPECIES OF Dalia FROM COLOMBIA AND Ecuapor (Hesrermpar) Stephen R. Steinhauser 53 


THE COLLAPSED EGGS FOUND IN THE BURSA COPULATRIX OF A PLUM MOTH, ILLIBERIS ROTUNDATA JORDAN 
(ZYGAENIDAE: PROGRIDINAE): AN UNUSUAL EGG RESORPTION SYSTEM? Chiharu  Koshio, — 


Yoshihisa Tanaka and: Shin-ichi Kudo-2--------- 2-4-2505 ee 
STUDIES IN THE GENUS HyLepHitA BiuLBEeRG, I. THe BOULLETI SPECIES GROUP (HESPERIIDAE: Hes- ice eee 

PERINAE) C. Don MacNeill -------------------------------------------9 nn nnn 605570 
DEW-DRINKING BY MALE MONARCH BUTTERFLIES, Danaus pLexippus (L.) Dennis Frey, Robert © ie As 

Roman and Lindsay Messett -----------------------------------------------------------------------= 90) eee 


SIBLING RIVALRY IN FLORIDA: THE DISPLACEMENT OF PyRGUS COMMUNIS BY PYRGUS ALBESCENS (HEsPERI- Goa 


IDAE) John V. Calhoun-------------------------------- =n noon nnn enn UgR ae 


GENERAL . Norms at 
IMMATURE STAGES OF Sars ROSALIA (NYMPHALIDAE, IrHomunaE) André V. L. Freitas and = 
Keith 8. Brown Jr, -----------------------------=-----~---$-- $n nn nnn nn nnn nnn nnn nnn nnn LOA ce 
MATINGS WITHOUT SPERMATOPHORE TRANSFER AND WITH TRANSFER OF TWO SPERMATOPHORES IN a ane 
Cattopurys XAMI (LycAENIDAE) Carlos Cordero -------------------------------------------- 106 
PAPILIO DEMOLEUS (PAPILIONIDAE) IN BORNEO AND Batt Kazuma Matsumoto------------------ LOSS ate 


Notes On THE NEVADA BUCK MOTH, HEMILEUCA NEVADENSIS (SATURNIIDAE) IN SOUTHERN MANI- 


mona DG: Henne and J. K Diehl a 12 . a 


This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). 


“Set Ab FY op Volume 56 Number 3 
er a | 11 October 2002 


ISSN 0024-0966 


Journal of the 
Lepidopterists Society 


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OC) BF Aue 


LIBRARIES 


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JOURNAL OF 


Tue LepiporrTerists’ SOCIETY 


Volume 56 


Journal of the Lepidopterists’ Society 
56(3), 2002, 117-122 


Number 3 


IMMATURES, NATURAL HISTORY AND THE SYSTEMATIC POSITION 
OF BIA ACTORION (NYMPHALIDAE) 


ANDRE V. L. FREITAS 


Museu de Histéria Natural and Departamento de Zoologia, Instituto de Biologia, 
Universidade Estadual de Campinas, CP 6109, Campinas, Sao Paulo, 13083-970, Brazil 


DEBRA Murray! 


Department of Entomology, 402 Life Sciences Bldg., Louisiana State University, Baton Rouge, Louisiana 70803, USA 


AND 


K. S. BROWN JR. 


Museu de Hist6éria Natural and Departamento de Zoologia, Instituto de Biologia, 
Universidade Estadual de Campinas, CP 6109, Campinas, Sao Paulo, 13083-970, Brazil 


ABSTRACT. The early stages of the Amazonian nymphalid butterfly Bia actorion were studied in four localities (three in Brazil and one in 
Ecuador). The eggs bore multiple transverse and longitudinal ridges. First instar larvae had a setose head-capsule without scoli. All other instars 


were characterized by basally fused, highly ramified and setose head scoli, and a bifurcate plate on the tenth abdominal segment. These ch 


ar- 


acters indicate that this species (and its monotypic tribe) belong within the Brassolinae. 


Additional key words: 


The genus Bia Hiibner is monotypic, and its only 
species, Bia actorion (Linnaeus, 1763), is found 
throughout the Amazon from low to mid elevations in 
the understory of dense forests (Masters 1970). Imma- 
tures of this species were unknown, and the only stud- 
ies on Bia were based on morphological characters of 
the adults (Miller 1968, Vane-Wright 1972). 

There is disagreement among authors regarding the 
systematic position of the genus Bia. Weymer (1912) 
placed it in the Satyridae, and his classification was 
maintained by Ehrlich (1958), Miller (1968) and Har- 
vey (1991); while Clark (1947, 1948) classified Bia in 
the Brassolidae, an opinion that was not widely fol- 
lowed. The currently accepted position of Bia follows 
Miller (1968) who included it in its own tribe Biini 
(subfamily Biinae, Satyridae), along with two other 


' Current address: Dept. of Entomology, Oregon State University, 


Corvallis, Oregon 97331-2907, USA 


Arecaceae, Brassolinae, life history, Satyrinae. 


tribes, Antirrhini and Melanitini. Based on a cladistic 
analysis of early stage characters, DeVries et al. (1985) 
revised Miller's (1968) classification of Biinae by trans- 
ferring Antirrhini to the Morphinae. Although Bia was 
not included in their analysis, DeVries et al. (1985) 
noted that it possesses exposed dorsal androconial 
patches similar to those of Caligo (Brassolinae), and 
suggested that early stages would be informative for 
determining the systematic position of Bia. 

This paper describes the immature stages of Bia ac- 
torion and discusses the systematic position of this 
species within the Nymphalidae. 


STUDY SITES 


Adults and immatures of Bia actorion were ob- 
served and collected in the field in four different local- 
ities in the Neotropics: the DBFF project reserves 
north of Manaus, Amazonas, Brazil (1981): Reserva 
Extrativista do Alto Juruaé, Upper Jurua River, 


118 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 1. Early stages and adult natural history of Bia actorion. a, egg; b, first instar; c, head capsule of first instar; d, lateral view of head cap- 
sule of first instar showing stemmata; e, antenna of first instar; f, second instar in molt; g third instar; h, fourth instar; i, j, fifth instar; k, frontal 
view of fifth instar head; I, m, pupa (ventral, lateral); n, perched adult flashing wings (dark brown, with blue and yellow on forewing) 


VOLUME 56, NUMBER 3 


119 


T1 T2-T3 A1-A2 A3-A6 A7 


Marechal Thaumaturgo, Acre, Brazil (1989-1998); 
Jatun Sacha Biological Station, upper Napo River, 
eastern Ecuador (1993), and Cristalino Jungle Lodge, 
Rio Cristalino, Alta Floresta, northern Mato Grosso, 
Brazil (2000). 


MATERIALS AND METHODS 


Immatures were collected in the field in Ecuador, 
while in Brazil eggs were obtained from a female con- 
fined in a plastic bag with leaves of the host plant (Alta 
Floresta, following Freitas 1991) or through expres- 
sion of the abdomen of gravid females (see Brown & 
Benson 1974). Immatures were reared in plastic con- 
tainers, cleaned daily, and fresh plant material was pro- 
vided every 2-3 days. Data were taken on behavior 
and development times for all stages, and head cap- 
sules and pupal castings were preserved. Immatures 
were fixed in Kahle (AVLF collection) or preserved in 
70% ethanol after being immersed in boiling water for 
several seconds (DM collection). The cephalic capsule 
was sputter coated with gold and observed in scanning 
electron microscopy (Jeol JSM-5800LV). Nomencla- 
ture of body setae follows Nakanishi (1988). 


RESULTS 


Host plants. The host plants of Bia actorion were 
palms (Arecaceae) in all study sites. In Ecuador the 
host plant was the spiny palm Astrocaryum murumuru 
Martius, 1824. In Alta Floresta females were observed 
flying around a species of Geonoma palm without 
spines, which was used successfully as host plant. In 
Manaus and the Upper Jurua the palm species were 
not identified, but were similar to, and possibly also 
members of the genus Geonoma, used for rearing the 


AQ A10 


Fic. 2. _Chaetotaxy of the first instar larva of Bia actorion. 


larva in the former case (transplanted in Monjolinho 
arboretum, Campinas, Sao Paulo, Brazil). 

Oviposition and larval behavior. At midday on 
May 2, 1993, at Jatun Sacha, Ecuador, a female was 
observed ovipositing on a A. murumuru palm. Eggs 
were laid singly on the spines emerging from new 
palm fronds, or on detritus caught on the palm fronds. 
Oviposition was a lengthy process, with the female 
resting for several minutes between each egg-laying 
episode. Once an oviposition site was selected, the fe- 
male rubbed her abdomen back and forth several 
times before laying eggs. The observed female laid six 
eggs on a single palm tree before moving off rapidly 
into the forest. 

In Ecuador, the host plant was a tall palm tree 
(about 15 meters high) while in Manaus the probable 
host was a very small plant. In Alta Floresta and Acre, 
females were also observed flying around small under- 
story palms. 

Once hatched, larvae were active and moved around 
on the hostplant, unlike the sluggish behavior of typi- 
cal satyrines (pers. obs.). 

Diagnosis for mature larva. Head dark brown 
with a pale brown stripe outlining frons and a pale 
brown lateral stripe; dorsal head scoli fused at base, 
ramified and with many setae; apical and subapical se- 
tae inserted into elongated tubercles. Body slender, 
green, with a bright white dorsal stripe; long bifid tail 
present on the last abdominal segment. 

Description of early stages. All life stages are de- 
scribed below based on specimens from Brazil and 
Ecuador. 

Egg (Fig. la). Gray-white when first laid, becoming 
purple and pinkish-gray as the developing larva takes 


form; spherical with 25-30 longitudinal ribs and up to 
50 less well developed transverse ribs; height 1.2 mm, 
diameter 1.4 mm (n = 10, Alta Floresta). Time of de- 
velopment: Ecuador—9 days (n = 6); Alta Floresta— 
8-10 days (n = 30). 

First instar (Fig. 1b—-e). Head brown; setae numer- 
ous, long, black, and plumose; setae projecting anteri- 
orly; two pairs of chalazae on vertex of head capsule. 
Head capsule width 0.96 mm; setae length 0.1-0.5 
mm. (material from Alta Floresta, Fig. lc). Body 
green, smooth, with one dorsal and three lateral red 
stripes, wider dorsally; T1 with a heavily sclerotized, 
smooth, prothoracic shield; more lightly sclerotized 
patch with cuticular spinules surrounding XD2 and ex- 
tending dorsolaterally to posterior edge of segment; 
XD and D setae on T1 with bulbous apex; excepting 
SV and V, all abdominal setae dark, heavily sclerotized 
at base and with sharp spines along shaft; SD1 filiform 
and long on thorax and AQ; L2 filiform but not long on 
thorax; caudal filaments well developed; D2 setae on 
A10 long. Body chaetotaxy illustrated in Fig. 2. Maxi- 
mum length 9 mm (Alta Floresta). Duration of instar: 
Ecuador—11 days for 4 larvae; Alta Floresta—8 days 
for 30 larvae. 

Second instar (Fig. 1f). Head dark brown dorsally 
and light brown ventrally and basally; two dark brown 
scoli at vertex of head capsule, fused into one trunk for 
dorsally projecting basal portion, then separating into 
two projecting dorsolaterally, with apex curving pos- 
terolaterally; setae on scoli plumose; two smaller pairs 
of dark brown subdorsal scoli on head capsule; ante- 
rior scolus projecting anteriorly and posterior scolus 
projecting posterolaterally, both scoli curved before 
apex; setae on head capsule fine and short, except se- 
tae surrounding stemmata and on base of head cap- 
sule, that are dark, thick, and plumose; microgranula- 
tions on surface of scoli and (more finely) on surface of 
head capsule; head capsule with weakly developed 
ridges in the integument. Head capsule width 1.1 mm 
(only 15% wider than first instar, but twice as high due 
to scoli); anterior scolus 0.6 mm; posterior scolus 0.8 
mm; scolus on vertex: fused trunk 0.6 mm, free arms 
1.1 mm (material from Alta Floresta). Body slender, 
dark green; dorsal median stripe bright white; red spot 
on A3 along midline; laterally reddish green from A8 
to posterior end; caudal projections long, dark green, 
and projecting laterally; numerous small secondary se- 
tae on body; SD1 filiform on T1-T3, A3-A7, and AQ; 
L2 also filiform on T1; all filiform setae with small, 
slightly elevated bases and light sclerotized patch. 
Maximum length 15 mm (Alta Floresta). Duration of 
instar: Ecuador—9 days (n = 3); Alta Floresta—14 
days (n = 25). 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Third instar (Fig. 1g). Head as in previous instar 
except for the following: head capsule all dark brown 
including basally, with pale brown stripe outlining 
frons and pale brown lateral stripe; setal bases sur- 
rounding stemmata and on base of head capsule larger, 
with setae emerging subapically; large, elongate setal 
bases present on apex of each scolus and several along 
shafts, all emerging subapically with apex smooth and 
pointed; integumental ridges outlining faint, shallow 
depressions on head capsule. Head capsule width 1.5 
mm; anterior scolus 0.8 mm; posterior scolus 1.2 mm; 
scolus on vertex: fused trunk 1.0 mm, free arms 1.8 
mm (material from Alta Floresta). Body same as in 
previous instar except lateral red coloration present on 
abdominal segments. Maximum length 30 mm (Alta 
Floresta). Duration of instar: Ecuador—10-11 days (n 
= 2); Alta Floresta—11 days (n = 20). 

Fourth instar (Fig. 1h). Very similar to third instar, 
with the general patterns darker than in previous in- 
stars. Head capsule width 2.1 mm; anterior scolus 1.3 
mm; posterior scolus 1.8 mm; scolus on vertex: fused 
trunk 1.8 mm, free arms 2.7 mm (material from Alta 
Floresta). Maximum length 35 mm (Alta Floresta). 
Duration of instar: Ecuador—9 days (n = 2); Alta Flo- 
resta—I1 days (n = 16). 

Fifth instar (Fig. li-k). Head same as previous in- 
stars except for the following: head capsule and scoli 
black frontally and brown basally, with a white stripe 
laterally on head capsule; scoli larger than previous in- 
star; more secondary setal bases on scoli and around 
stemmata and base of head capsule forming elongate 
projections with subapical setae; setal bases of these 
setae also larger than in previous instar. Head capsule 
width 2.9 mm; anterior scolus 2.2 mm; posterior scolus 
2.5 mm; scolus on vertex: fused trunk 2.5 mm, free 
arms 3.1 mm (Alta Floresta). Body slender, green; dor- 
sal median stripe bright white; stripe narrows abruptly 
at anterior edge of each segment and then widens to 
posterior edge, with size change greatest on Al to A6; 
within white stripe thin gray line along midline and 
reddish gray coloration, darkest on A3, lateral to gray 
line at widest portion of white stripe; small white dots 
dorsolaterally and reddish-brown coloration ventrolat- 
erally; caudal filaments long, dark greenish-gray, pro- 
jecting laterally; secondary setae on body dark brown, 
wide, and flattened; filiform setae on body with heavily 
sclerotized and small setal bases, projecting only mini- 
mally from body surface and surrounded by large, 
sclerotized patch. Maximum length 55 mm (Alta Flo- 
resta). Duration of instar: Ecuador—10 days (n = 2); 
Alta Floresta—14 days (n = 13). 

The larvae from Ecuador passed through an addi- 
tional sixth instar. The head capsule was the same as in 


VOLUME 56, NUMBER 3 


the previous (fifth) instar; the body was the same ex- 
cept for more extensive reddish-gray coloration within 
the white stripe, reducing white coloration to a thin 
outline, and white dots dorsolaterally larger. Duration 
of instar: 22-25 days (n = 2). 

Pupa (Fig. 1 1-m). Short and bumpy, slightly curved 
anteriorly and strongly curved at A4. Entirely green, 
with dark lateral keels and a shiny dot near the wing 
caps. Total length 15 mm (Alta Floresta). Duration of 
pupa: Alta Floresta—17 days (n = 8). 

Adult behavior. Adults were found in dense forest 
with abundant small palms, including gallery (riparian) 
forests in an upland cerrado (savannah) landscape (Vil- 
hena, 600 m, SE Rondénia, near Mato Grosso border, 
Brazil). 

Adults fly in the understory and are active from be- 
fore dawn to nightfall, being easily found in patches 
with rotting fruits on which they feed. During after- 
noons and on cloudy days, [the] males can be found in 
small openings in the vegetation, often near streams, 
but not always near areas with palms. Through mark- 
ing of individuals (DM) males were found to persist 
for up to 20 days. Although some individuals moved 
from patch to patch over time, most remained loyal to 
a particular area. Interactions between males were 
common. No mating attempts with passing females 
were observed, although courtship behaviors were 
commonly seen. Courtship appeared to be lengthy and 
complex, including several different tandem flight pat- 
terns, male contact with females during flight, and solo 
male flight over perched females while they flash their 
wings. Occasionally courtship behaviors were observed 
between two males. 


DISCUSSION 


Regarding the controversy about the classification of 
Bia (Vane-Wright 1972, DeVries et al. 1985), the early 
stages give additional clues about its possible system- 
atic position. The eggs, with many transverse and lon- 
gitudinal ridges, and the first instar larvae with the 
head capsule lacking any kind of scoli and bearing long 
setae, giving a furry appearance, are most similar to 
those of the Brassolinae (Casagrande 1979, 1992, De- 
Vries 1987). Although some Satyrinae have eggs with 
ridges (Emmel & Mattoon 1972), eggs of Brassolinae 
are distinctive in having many well-marked ridges 
(30-60), considered a trait exclusive to this subfamily 
(Freitas 1999). The mature larva also bears a typical 
brassoline feature, three pairs of scoli on the head cap- 
sule. This condition is observed in most Brassolinae 
(except in the gregarious Brassolis, that lack the head 
projections) and is absent in all known satyrine larvae, 
that show head capsules bearing a single pair of scoli 


on the vertex (pers. obs). Additionally, palms are com- 
mon host plants of Brassolinae, whereas in Satyrinae 
palm feeding is rare (DeVries 1987, Ackery 1988). 

Based on adult morphology, Bia was suggested to be 
more closely related to Brassolinae than to Satyrinae 
(Vane-Wright 1972, DeVries et al. 1985). Our exami- 
nation of early stages strongly supports this hypothesis. 
Confirmation must await a thorough phylogenetic 
study and broader sampling of immatures among 
Satyrinae and Brassolinae. 


ACKNOWLEDGEMENTS 


AVLF and KSB are very grateful to the personnel of the Floresta 
Amazonica Hotel in Alta Floresta, MT, and the Rio Cristalino Jungle 
Lodge (Vit6ria de Riva Carvalho and her son Edison Carvalho), for 
housing and food, extensive and patient logistic support, during 
work in this region in February and June 2000. The ASAREAJ pro- 
vided logistic support during fieldwork in Acre, and the PDBFF 
supported work in the reserves north of Manaus. DM is grateful to 
Alejandro Suarez for his encouragement and help in rearing the lar- 
vae, and to the United States Peace Corps for their support of this 
research. We would like to thank Drs. Carla M. Penz, David Lees 
and P. J. DeVries for comments on the manuscript. Karina C. 
Mancini helped with the SEM methods. Financial support for 
AVLF and KSB was provided by [the Brazilian] CNPq, and by 
grants from the BSP and CIFOR in Acre. This paper was partially 
finadledl by the BIOTA/FAPESP program, grants 98/05101-8 and 
00/01484-1. 


LITERATURE CITED 


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mata (Lepid. Nymph.). Biotropica 6:205-208. 

CASAGRANDE, M. M. 1979. Sobre Caligo beltrao (Illiger). I. Tax- 
onomia, biologia, morfologia das fases imaturas e distribuig6es 
espacial e temporal (Lepidoptera, Satyridae, Brassolinae), 
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. 1992. Narope cyllene Felder & Felder, sp. rev. Re- 
descrigao e fases imaturas (Lepidoptera, Nymphalidae, Bras- 
solinae). Revta. Bras. Zool. 7:129-145. 

Ciark, A. H. 1947. The interrelationships of the several groups 
within the butterfly superfamily Nymphaloidea. Proc. Ent. Soc. 
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1948. Classification of the butterflies, with the allocation of 
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DEVniEs, P. J. 1987. The butterflies of Costa Rica and their natural 
history. Papilionidae, Pieridae, Nymphalidae. Princeton Uni- 
versity Press, Princeton, New Jersey. 

DeVries, P. J., I. J. Kircuinc, & R. I. VANE-Wricut. 1985. The 
systematic position of Antirrhea and Caerois, with comments 
on the higher classification of the Nymphalidae ( Lepidoptera). 
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EHRLICH, P. R. 1958. The comparative morphology, phylogeny and 
classification of the butterflies (Lepidoptera: Papilionoidea). 
Univ. Kansas Science Bull. 39:305—370. 

EMMEL, T. C. & S. O. Matroon. 1972. Cercyonis pegala blanca, a 
“missing type” in the evolution of the genus Cercyonis (Satyri- 
dae). J. Lep. Soc. 26:140-149. 

Freitas, A. V. L. 1991. Variagéo morfolégica, ciclo de vida e sis- 
tematica de Tegosa claudina (Eschscholtz) (Lepidoptera, 
Nymphalidae, Melitaeinae) no Estado de Sao Paulo, Brasil. 
Rey. bras. Ent. 35:301—306. 

. 1999. Nymphalidae (Lepidoptera), filogenia com base em 


122 


caracteres de imaturos, com experimentos de troca de plantas 
hospedeiras. PhD. Thesis, Universidade Estadual de Campinas, 
Campinas, Sao Paulo. xii + 170 pp. 

Harvey, D. J. 1991. Higher classification of the Nymphalidae. Ap- 
pendix B, pp. 255-273. In Nijhout, H. F. The development and 
evolution of butterfly wing color patterns. Smithsonian Institu- 
tion Press, Washington, D. C. 

Masters, J. H. 1970. Bionomic notes on Haeterini and Biini in 
Venezuela (Satyridae). J. Lepid. Soc. 24:15-18. 

MILLER, L. D. 1968. The higher classification, phylogeny and zoo- 
geography of the Satyridae (Lepidoptera). Mem. Am. Entomol. 
Soc. 24:iii + 174 pp. 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


NAKANISHI, A. 1988. Study on the first instar larvae of the Subfam- 
ily Nymphalinae (Lepidoptera, Nymphalidae). Spec. Bull. Lep. 
Soc. Jap. 6:83-99. 

VANE-WRIGHT, R. I. 1972. Pre-courtship activity and a new scent 
organ in butterflies. Nature 239:338-340. 

WEYMER, G. 1910-1912. 4. Family Satyridae. pp. 173-280. In A. 
Seitz (ed.), Die Grossschmetterlinge der Erde. Stuttgart, A. 
Kernen. 5 (Die Amerikanische Tagfilter). 


Received for publication 25 August 2001; revised and accepted 12 
December 2001. 


Journal of the Lepidopterists’ Society 
56(3), 2002, 123-128 


BIOLOGY OF ANTHERAEA ANDAMANA (SATURNIIDAE) 
ON THE ANDAMAN ISLANDS, INDIAN OCEAN 


PRASHANTH MOHANRAJ AND K. VEENAKUMARI 
Central Agricultural Research Institute, PB. No. 181, Port Blair, Andaman Islands, India! 


ABSTRACT. Antheraea andamana was collected from the forests of South Andaman and Little Andaman, two islands in the Bay of Ben- 
gal, Indian Ocean. This is the first time that a saturniid is being reported from an island other than S. Andaman in this archipelago. The imma- 


ture stages are described and illustrated for the first time. 


Additional key words: Andamans, immatures, Ficus. 


Antheraea Hiibner (1819) is the largest saturniid 
genus in the Oriental Region, with species occurring 
from northeastern India to the western Moluccas and 
the eastern portion of Palaearctic Asia (Nassig et al. 
1996a). In the Andaman islands this genus is repre- 
sented by just one endemic species, Antheraea an- 
damana Moore (Moore 1877, Watson 1911, Peigler 
1989) (Fig. 1). Along with Actias ignescens Moore (an- 
other endemic species) and Actias selene (Hiibner 
1806) (now A. callandra Jordan, 1911; see Prashanth 
Mohanraj et al. 1996), it was among the first saturniids 
to be reported by Moore (1877) from the Andaman 
and Nicobar islands. 

Here we report on the first captures of the imma- 
ture stages of A. andamana from South and Little An- 
damans, and provide information on the natural his- 
tory of this poorly known insular taxon. 


METHODS 


Both adults and immatures were collected from 
South Andaman and Little Andaman. The two islands 
are separated by the 46 km wide Duncan Passage in 
the Bay of Bengal, Indian Ocean. 

On South Andaman all stages of the moth were 
collected from Ficus trees, at three sites: (1) in the 
evergreen forest at Mount Harriet, (2) along the 
roadside at Homfraygunj where the original forest 
has been cleared and is now being cultivated, and (3) 
in the back mangals at Chiriyatapu. Mt. Harriet rises 
from sea level to a height of 365 m. A large Ficus 
arnottiana Migq. grows at the summit, from which lar- 
vae of A. andamana were collected. Eggs, larvae and 
pupae were also collected from Ficus altissima 
Blume at lower elevations along the Mt. Harriet 
range. A few F. altissima trees growing on roadsides 
in Homfraygunj yielded eggs, larvae and cocoons of 
A. andamana. Larvae and cocoons were taken from a 
group of F. Pretusa L. trees in a stand of mangroves 
which had their bases periodically submerged by the 
rising tide. 


‘Current address: Project Directorate of Biological Control, P.B. 


No. 2491, H.A. Farm Post, Hebbal, Bangalore- 560 024, India. 


At Little Andaman moths, eggs, larvae and cocoons 
were collected from the foliage of a stand of Ficus 
Pretusa growing in a swampy area (Fig. 2) close to the 
sea. 

All the material was collected from their host plants 
in forests and clearings. Several collecting methods 
were used. First, groups of 4 to 6 people inspected 
the foliage of trees manually with a pair of binoculars. 
Second, the ground beneath the canopy was searched 
for fecal pellets. Since the larvae apparently do not 
move large distances during their larval period the 
presence of feces, if fairly fresh, is a clear indication of 
their presence. Third, a long nylon rope with a stone 
tied to one end was thrown over high branches, and 
these were pulled down and scanned leaf by leaf. 
Lastly, trees were climbed and their foliage searched 
manually, 

The largest number of A. andamana individuals was 
obtained by the fecal pellet method. This was also the 
most efficient in terms of number of individuals ob- 
tained per unit of search time. The primary disadvan- 
tage of this method was that only larvae were located. 
It also yielded a greater number of older than younger 
instars, as the larger fecal pellets of the later instars are 
more easily spotted. 

All material collected was brought to the laboratory 
in individual plastic containers of various sizes (in 
keeping with the size of the stage caught). Eggs and 
larvae were then transferred onto bouquets of their 
food plants in flasks of water whose openings were 
tightly secured with cotton plugs. Each larva was then 
placed in a separate cage. Observations were made 
daily in the morning when the cages were cleaned and 
larvae provided with fresh food. 


NATURAL HISTORY 


From our rearing experiences, the following de- 
scriptions of the immature stages of A. andamana can 
be offered (Table 1 and Fig. 3): 

Egg. Dorsoventrally flattened with an irregular 
brown patch on top. Chorion mottled, mostly brown 
with small irregular dirty white patches. Chorionic sur- 
face honey-combed with shallow depressions. No 


Fic. 1. 


Adult females of Antheraea andamana Moore. 


brown bands encircling the eggs. Inner surface pale 
brown, smooth and highly reflective. The eggs were 
laid singly or in batches of 2 or 3 on the ventral sur- 
faces of leaves. Some were laid about 1 cm apart while 
others were laid in contact with each other. 


TABLE 1. 
Time of 
development (days) Egg 1 2 
All individuals 8.7+0.8 3.9 + 0.6 3.5 + 0.5 
im = 12 me Ie im = 17 
Males only = 3.8 + 3.6 + 0.6 
5 n=5 
Females only = 3.8 + 0.5 3.6 + 0.6 
n=9 n=9 
Dimensions (cm) 
All individuals 3.7 + 0.2! 0.7+0.1 16+0.1 
X n= 30 n= 18 
3.3 + 0.2 
in = 30) 
Males only = 0.7+0.1 1.5+0.1 
n=9 n=9 
Females only = 0.7 1.7+0.1 
n=4 m= 
Head capsule width (mm) 1.8 +0.1 2.9+0.1 
in = 7 n = 20 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 2. Brackish water swamp habitat of A. andamana at Little 
Andaman. A freshly emerged female moth was collected from the 
over hanging branches of Ficus Pretusa on the left of the picture. 


First instar. Head deep brown to black, glossy 
with sparse white setae; clypeus, labrum and labium 
white; antennae/palp semitransparent golden brown 
or honey colored arising from a whitish base. Protho- 
racic shield glossy and of the same color as the head. 
Thoracic and abdominal segments creamish-yellow 
dorsally, tending towards white or pale yellow ven- 
trally. A narrow brown band or line is present towards 
the anterior and posterior margins of each segment. 
The scoli are located between these bands. These 
bands take on a deeper hue a little above the lateral 


Developmental time and dimensions of the immature stages of Antheraea andamana on the Andaman islands. 


Larval instars 


3 4 5 Pupa Total 
4.2+0.8 5.9414 9.1+0.9 25.1 + 3.0 60.7 
n = 20 n = 20 n=17 n=13 
3.6 +0.9 5.2 + 0.5 8.4+0.9 25.0 + 1.4 49.5* 
n= 9 n=9 1 = &) n=4 
46+0.6 5.2 + 0.5 9.0 27.4+2.7 53.6* 
in = 5) im = 5) n=5 m=) 
2.8+0.2 3.8 + 0.2 5.2 +0.4 DH) a= OLSk — 
m= 20) n = 20 ml = II) x 
2340.3 
n=3 
2.9 + 0.2 3.7 + 0.2 5.0 + 0.2 = 
n= o) im = 15) n= 
2.8+0.3 3.7 + 0.2 5.4 + 0.3 - = 
m = 4! n=4 n=4 
4.1+0.1 5.9 + 0.2 = = = 
n= 21 m = il 


‘Measurements in mm. 
xclusive of incubation period. 
Dimensions of cocoon, not of pupa. 


VOLUME 56, NUMBER 3 125 


Fic. 3. Immature stages of Antheraea andamana Moore. a, Eggs. b, First instar larva. e, Second instar larva. d, Fifth instar larva. e, Fifth 
instar larva—close up of head. f, Fifth instar larva—anal proleg. g, Cocoon. h, pupa in ventral view. 


126 


scoli on the first to seventh abdominal segments 
while they retain the same intensity of brown 
throughout their length on the remaining segments. 
All scoli, with the exception of the dorsal scoli on the 
prothorax, basally yellow (of the same color as the 
rest of the body); terminally bulbous and transparent 
yellow from which arise five transparent, whitish se- 
tae tapering, towards their terminal ends and 
arranged in a rough circle from the center of which 
arises another seta. The dorsal scoli on the prothorax 
are very short and appear to have two conjoined 
heads with eight long whitish setae. Single, dorsal 
scolus on the eighth abdominal segment. Anal plate 
and outer lateral surface of anal prolegs deep brown. 
Spiracles on prothorax internally of the same color as 
the rest of the body with a narrow brown rim. Ab- 
dominal spiracles also brown rimmed, but with 
brown striations arising from the rim and running to- 
wards the center. Legs black with sparse, short, white 
setae and brown claws. Prolegs black distally with 
nine transparent white setae in the black region and 
brown crochets. 

Second instar. Head deep brownish-black, with 
thin black sparsely distributed setae. Frons and outer 
margins of frontal and epicranial sutures pale brown; 
antennae brown, set in a white base; maxillary palp 
and labium partly white and partly black; labrum 
golden brown; clypeus white. Thoracic and abdomi- 
nal segments yellow-green with small white, club- 
shaped setae. Prothracic shield brown in the center 
of the anterior margin, becoming progressively paler 
laterally and posteriorly. Dorsal scoli on prothorax 
short, situated more towards anterior margin and 
with about 7 to 8 setae. The largest scoli are the dor- 
sal scoli on the meso- and meta-thoracic segments. 
They are also the only tri-colored scoli being black 
terminally, followed by an orange-brown band and 
yellow basally. All other dorsal scoli are small with an 
orange brown ring/band. On the eighth abdominal 
segment the dorsal scoli are fused but distinctly bi- 
furcated on top with § brownish setae (6 forming a 
rough circle with 2 in the center). Subdorsal and sub- 
spiracular scoli are small, orange-brown with termi- 
nal crown of black or brown setae. Prothoracic spira- 
cle yellowish; all other spiracles deep brown to black 
except for that on the eighth abdominal segment 
which is brownish-white. Subspiracular line faint yel- 
low anteriorly and more prominent posteriorly par- 
ticularly on segments 7 and 8 where it gradually turns 
brown. It is narrow anteriorly, becoming broader to- 
wards the rear. The anal plate is black and fringed 
with brown setae along the outer margins. The lateral 
surface of the anal prolegs have a deep brown to 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


black triangle with a basal yellow patch. Ventral sur- 
face is yellowish-green with short white and longer 
brown fine setae on first and second abdominal seg- 
ments. Legs are black with brown claws while the 
prolegs are yellowish brown distally with black setae 
arising from the center of small, black, circular spots. 
Crochets are brown. 

Third instar. Head brown to black in color with 
white clypeus and white and brown labrum, Bases of 
antennae a rich cream to yellow. The rest of the larva 
is greenish yellow. Prothoracic shield yellow, tinged 
with brown anteriorly. Scoli very small and brown 
with one long and 5 to 6 short, spiny, brown setae. 
Dorsal scoli on meso and meta thorax terminally deep 
brown to black with an annular brown ring beneath 
them and the basal two-thirds distinctly yellow (more 
intense than the rest of the body). The subspiracular 
scoli in this instar are blue on all the segments, the 
blueness being most pronounced on the first three ab- 
dominal segments. Spiracles on the first seven ab- 
dominal segments are black while that on the eighth 
abdominal segment is pale brown with a silvery-white 
longitudinal line in the center. The lateral yellow line 
is prominent. It is darker and broader towards the 
anal end becoming progressively narrower and less 
distinct as it approaches the first abdominal segment 
where it terminates. Though greenish-yellow overall, 
the “green-ness’ is more pronounced below the lateral 
line than above it. Ventrally, all the thoracic segments 
and the posterior abdominal segments from seven to 
the anal tip are yellow. The remainder of the ventral 
surface is green with a mid-ventral yellow line. Legs 
deep brown to black with brown claws. Prolegs 
distally brown and yellow proximally. In the yellow 
area are circular brown spots of variable size and each 
has a brown seta in the center-most are long, a few are 
short. Laterally, the anal prolegs have a brown 
triangle. 

Fourth instar. Head and mandibles deep burnt 
brown; frons and labrum cream, clouded with brown. 
The rest of the body (including the ventral surface) is 
pale green—of the same shade as the under surface of 
the leaf on which it feeds. (The dorsal surface of the 
leaf is a deeper green). Short brown setae are sparsely 
distributed over the entire body surface. Prothoracic 
shield is cream anteriorly and brown posteriorly. The 
yellow supraspiracular line is broad posteriorly and 
narrows progressively towards the anterior end to ter- 
minate abruptly above the first abdominal spiracle. It 
does not extend onto the thoracic segments. Dorsal 
scoli on thorax not prominent but visible as pale yellow 
circular patches with brown centers. Spiracles deep 
brown and ringed with a narrow pale yellow band. 


VOLUME 56, NUMBER 3 


Fifth instar. Head burnt brown in color with 
labrum, clypeus, frons and broad regions on either 
side of the epicranial suture pale dirty brown. The rest 
of the larva is pale green, becoming a shade darker be- 
low the spiracles. Prothoracic shield also burnt brown. 
Supraspiracular line yellow and enclosing the top one- 
third of the eighth abdominal spiracle. This line passes 
above the seventh abdominal spiracle and keeps going 
progressively higher up to the first abdominal segment 
where it abruptly stops. The row of black setae on the 
lateral scoli below the spiracle on each abdominal seg- 
ment arises from a small, pale blue patch on segments 
one to six, with the last two patches being very small 
and very faintly blue. 

Cocoon. Stalked and spun between leaves such that 
it is almost totally covered with leaves. It is brown in 
color, greying as it ages; and the opening from which 
the adult emerges faces the petiole. 

Biology. Eggs, larvae and cocoons were found on Fi- 
cus altissima Blume, Ficus arnottiana and Ficus Pretusa 
(Moraceae). The larvae completed their life cycles 
when reared on the foliage of all these species of trees 
in the laboratory. We observed various stages of the 
moth on their host plants in the months of January, Feb- 
ruary, March, September, October, November and De- 
cember. Two species of hymenopterous parasitoids 
emerged from eggs collected at various times from the 
field: both species exhibited superparasitism with all in- 
dividuals of one species emerging from a single exit hole 
while all individuals of the second species emerged by 
making multiple exit holes in the chorion of the eggs. 
Some larvae succumbed to attack by the nuclear poly- 
hedrosis virus while being reared in the laboratory. 


DISCUSSION 


Moore (1877) described A. andamana from speci- 
mens in the Natural History Museum, London. He 
neither illustrated nor furnished characters to distin- 
guish it from other members of the genus. Cotes 
(1891-1893) was unable to verify its status as he could 
not access the specimens in London and there were no 
specimens in the Indian Museum (now the Zoological 
Survey of India, Calcutta). This led him to list this 
species along with four others as requiring confirma- 
tion of their specific status. It is also significant that 
Arora and Gupta (1979) do not even list it among the 
seven species of Antheraea that they deal with in their 
study of Indian non-mulberry silk moths. Apparently 
the Z.S.I. had failed to procure specimens from the 
Andaman islands in the 100 years since its first de- 
scription. 

It is pertinent to state here that since sericulture was 
not practised on the Andaman islands genetic contam- 


ination of the native saturniids is unlikely to have oc- 
curred by release of cultivated stock. 

Antheraea andamana belongs to the frithi subgroup 
of the paphia/frithi group of the subgenus Antheraea 
(Niissig et al. 1996b). So far the immatures of only a few 
species in this subgroup are known and, of these, most 
are only partially known (Nissig et al. 1996b, Holloway 
et al. 1996 and references therein). Only the life history 
of A. rumphii rumphii C. Felder from Ambon, Indone- 
sia has been fully worked out (Paukstadt et al. 1996). 

The only preimaginal character that Nassig (1991) 
and Niassig et al. (1996a) currently use in defining the 
paphia/frithi group is the usual presence of “a double 
ring of brownish color around the top and bottom of 
the flattened egg”; also known as “equatorial lines” 
(Jolly 1980). These lines are obsolete in A. andamana 
though present in its sister taxon A. rwmphii (Pauk- 
stadt et al. 1996). If this is found to be a useful charac- 
ter when the ova of more species in the frithi subgroup 
are described then perhaps the presence/absence of 
this character would serve to further subdivide this 
large and diverse subgroup. Nevertheless, until more 
information is generated on the biology and preimagi- 
nal morphology of species in the paphia/frithi group it 
will remain difficult to characterise the “limits of the 
different species” (Niassig et al. 1996a). 

As reported by Paukstadt et al. (1996), we also did 
not find silvery patches encircling the dorsal and 
supraspiracular scoli in any of the instars. Though Nis- 
sig et al. (1996b) say that the presence of these patches 
or rings is only facultative, they qualify their statement, 
saying that their total absence varies individually. We 
found no traces of these patches in over 30 larvae bred 
in the laboratory. Nassig et al. (1996b) state that the 
dorsal scoli on A8 are at least basally fused, even if not 
totally fused, in the known mature caterpillars. In ma- 
ture larvae of A. andamana these scoli on A8 maintain 
their distinct identities; it is only in the early instars 
that they are basally fused in A. andamana. 

Antheraea paphia L. is the only species in the genus 
Antheraea so far known to utilize species of Ficus as lar- 
val food plants (Stone 1991). Ficus spp. have now been 
found by us to be fed on by A. andamana, a species in 
the frithi subgroup, but in the same group as A. paphia. 
Antheraea moultoni Watson and Antheraea brunei 
Allen and Holloway are the only species in this genus 
that have been collected in or close to mangroves (Hol- 
loway 1987). The life histories of these species remain 
unknown, though first instars of A. brunei were ob- 
tained from eggs by Naumann (1994); these did not 
take any of the food plants he offered; Ficus was not in- 
cluded among them. Nissig et al. (1996a) however infer 
that A. brunei is a mangrove specialist. One of the habi- 


tats from which A. andamana has been collected by us 
is from brackish water Ficus swamps behind coastal 
mangroves in Little Andaman. It was also bred on this 
species of Ficus. These Ficus swamps are much drier in 
summer than during the monsoon, making them sea- 
sonal as opposed to permanent swamps. In South An- 
daman, on the other hand, we have found them breed- 
ing in the back mangals where the bases of the trees are 
submerged throughout the year during high tide. 

Rumphius (1627-1702) was the first to describe and 
illustrate the larva and cocoon of a species of Anther- 
aea (namely A. rumphii C. Felder) along with its larval 
food plant (Sonneratia caseolaris L.: Sonneratiaceae) 
(Diakonoff, 1959). Paukstadt et al. (1996), referring to 
the ‘Herbarium Amboinense’ misinterpreted the host 
plant ‘Mangium Caseolare Rubrum’ as Rhizophora 
caseolaris L. This error may also be found in Arora and 
Gupta (1979) and Stone (1991). “Mangium’ is a 
generic name that Rumphius applied to all mangrove 
plants (Tomlinson 1986). ‘M.C. Rubrum’ as mentioned 
and illustrated in ‘Herbarium Amboinense’ is S. caseo- 
laris L. (de Wit 1959). A. rumphii also belongs to the 
frithi subgroup like A. andamana. 

We note here that the Italian zoologist-cum-anthro- 
pologist Cipriani (1966) apparently had earlier made 
some observations on A. andamana (he called it A. pa- 
phia) when he was residing on Little Andaman studying 
the Onge, one of the four negrito tribes of these islands. 
He mentions Ficus as the larval food plant and describes 
how the larvae spin their stalked/petiolated cocoons. He 
says there are two broods, with the adults appearing in 
mid September and mid February. The adults, after 
emerging from their cocoons and spreading their wings, 
were observed by him to ‘sleep for two days, to wake up 
on the second night’ when the ‘active males seek and 
mate with the near sedentary females.’ He also noted 
that ‘virgin females attracted males from many kilome- 
ters away as is the case with Saturnia pyri.’ This is an ob- 
servation that we have so far not been able to repeat with 
any of the saturniids from the Andaman islands. 


ACKNOWLEDGMENTS 


We are grateful to A.K. Bandyopadhyay, Director, CARI, for en- 
couragement. We are indebted to W.A. Nissig and R.S. Peigler for 
‘atten us into the world of saturniids and for literature on this 
group of moths. Special thanks to J.D. Holloway for help in a myriad 
ways including comments on this paper. S. Naumann helped with 
literature and discussion. We thank B. Pardasaradhi for the pho- 
tographs and P.V. Sreekumar for identifying the plants. The assis- 
tance rendered by Om Prakash, Bikas Mondal, T. Das and Kinoo 
Ram in the field is gratefully acknowledged. We also thank the De- 
partment of Forests, Andaman and Nicobar Islands, for permission 
to collect insects from the National Parks and Wildlife Sanctuaries of 
these islands for scientific study. 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


LITERATURE CITED 


Arora, G. S. & I. J. Gupta. 1979. Taxonomic studies on some of 
the Indian non-mulberry silk moths (Lepidoptera: Saturniidae: 
Saturniinae). Mem. zool. Surv. India 16 (1):1-63. 

Cipriani, L. 1966. The Andaman Islanders. Frederick A. Praeger, 
Inc., New York. 159 pp. 

Cotes, E. C. 1891-1893. The wild silk insects of India. Indian 
Mus. Notes 2:69-89, pl. 2-15. 

Diakorr, A. 1959. Rumphius as an entomologist, pp. 127-144. In 
H. C. D. de Wit (ed.), Rumphius Memorial Volume. Uitgeverij 
en Drukkerij Hollandai N. V. — Baarn. 

Ho.ioway, J. D. 1987. The moths of Borneo [part 3]: superfamily 
Bombycoidea: families Lasiocampidae, Eupterotidae, Bomby- 
cidae, Brahmeidae, Satumiidae, Sphingidae. Kuala Lumpur 

(Southdene Sdn. Bhd.). 199 pp. + 163 figs. + 20 col. Pls. 

Houtoway, J. D., S. NAUMANN & W. A. Nassic. 1996. The Anther- 
aea Hiibner (Lepidoptera: Saturniidae) of Sulawesi, with de- 
scriptions of new species. Part 2: The species of the frithi- 
group Nachr. Entomol. Ver. Apollo, N.F. 17(3):225—-258. 

Jouty, M. S. 1980. Distribution and differentiation in Antheraea 
species (Saturniidae: Lepidoptera). [offprint of the paper pre- 
sented at the XVI Intemational Congress of Entomology, Ky- 
oto, Japan, 3-9 August, 1980]:1—20. 

Moore, F. 1877. The. lepidopterous fauna of the Andaman and 
Nicobar islands. Proc. zool. Soc. London 40:580-632. 

NAssic, W. A. 1991. New morphological aspects of Antheraea 
Hiibner and attempts towards a reclassification of the genus 
(Lepidoptera: Saturniidae). Wild Silk Moths ’89~90 (eds. H 
Akai and M. Kiuchi):1-8 

Nassic, W. A., R. E. J. LAMPE & S. Kacer. 1996a. The Saturniidae 
of Sumatra (Lepidoptera). ). Heterocera Sumatrana 10:1—110. 

1996b. Saturniidae of Sumatra. Appendix I: the preimagi- 
nal instars of some Sumatran and South East Asian species of 
Saturniidae, including general notes on the genus Antheraea 
(Lepidoptera). Heterocera Sumatrana 10:111—170. 

NAUMANN, S. 1994. Bemerkungen zu einer rotbraunen Form von 
Antheraea brunei Allen & Holloway (1985) Sowie zu den 
Priimaginalstadien der Art (Lepidoptera: Saturniidae). Nachr. 
entomol. Ver. Apollo, N. F. 15:359-362. 

PAUKSTADT, L. H., U. PAukKsTapT & S. NAUMANN. 1996. Die 
Praimaginalstadien von Antheraea rumphii rumphii C. Felder, 
1861 von Ambon, Indonesien, sowie taxonomische Ber- 
merkungen (Lepidoptera: Saturniidae) Entomol. Zeit. 
106(5): 165-176. 

PEIGLER, R. S. 1989. A revision of the Indo-Australian genus Atta- 
cus. Lepidoptera Research Foundation Inc., California. 167 pp. 

PRASHANTH MOHANRAJ, K. VEENAKUMARI, & R. S. PEIGLER. 1996 
[1993]. The host plant and pre-imaginal stages of Actias callan- 
dra (Lepidoptera: Saturniidae) from the Andaman Islands, Bay 
of Bengal, Indian Ocean. J. Res. Lepid. 32:16-25. 

STONE, S. E. 1991. Food plants of World Saturniidae. Mem. Lepid. 
Soc. 14:1-186. 

TOMLINSON, P. B. 1986. The botany of mangroves. Cambridge Uni- 
versity Press, London. 

Watson, J. H. 1911. The wild silk moths of the world, with special 
reference to the Saturnidae (sic). Delivered before the Textile 
Society, Manchester School of Technology, 4 Dec. 1911. 8 pp. 4 
col. pl. 

DE Wit, H.C. D. 1959. A checklist to Rumphius’ Herbarium Am- 
boinense, pp. 339-460. In H. C. D. de Wit (ed.), Rumphius 
Memorial Volume. Uitgeverij en Drukkerij Hollandai N.V. - 
Baarn. 


Received for publication 20 September 2000; revised and ac- 
cepted 23 November 2001. 


Journdl of the Lepidopterists’ Society 
56(3), 2002, 129-150 


A REVISION OF THE GENUS BULIA WALKER (LEPIDOPTERA: NOCTUIDAE) 


MICHAEL G. POGUE 


Systematic Entomology Laboratory, PSI, Agricultural Research Service, U.S. Department of Agriculture, c/o National Museum of Natural 
History, Smithsonian Institution, Washington, DC 20560-0168, USA, E-mail: mpogue@sel.barc.usda.gov 


AND 


A. CAROLINA LAUGHLIN 
103 N. Cuates, Los Fresnos, Texas 78566, USA 


ABSTRACT. A phylogenetic revision of the genus Bulia Walker is provided and includes a key to species, descriptions, illustrations of 
adults, both male and female genitalia, and distribution maps. The relationships among the five species of Bulia are based on an analysis of 16 
characters. The monophyly of Bulia is suported by 7 nonhomoplasious synapomorphies. Arsisca bolinalis Walker 1866 is proposed as a new syn- 
onym of Bulia confirmans (Walker) 1858 and Bulia morelosa Richards 1941 as a new synonym of Bulia similaris Richards 1936. Larval host 
plants are recorded for the first time for Bulia confirmans, on Jatropha gossypiifolia (L.) (Euphorbiaceae), and for Bulia mexicana (Behr), on 


Prosopis juliflora (Sw.) DC. (Fabaceae). 
Additional key words: 


The genus Bulia Walker 1858 was first revised by 
Richards (1936a). In this revision, he described a new 
species, B. similaris Richards, designated a neotype for 
B. mexicana (Behr) 1870, and concluded that these two 
species can only be separated from B. deducta (Morri- 
son) 1875 by structures of the male and female geni- 
talia. The maculation of the adult is useless due to the 
tremendous amount of inter- and intraspecific variabil- 
ity and degrees of sexual dimorphism that these species 
exhibit. Richards (1936b) described and diagnosed Bu- 
lia in regards to the Phoberia-Melipotis-Drasteria-Bo- 
ryza group of catocaline genera. A second revision by 
Richards (1941) incorporated additional material from 
Mexico and Central America and included two new 
species, B. schausi and B. morelosa, and a race of B. 
similaris from California (californica). 

The need for this third revision came about during 
the curation of the Nearctic Catocalinae at the Na- 
tional Museum of Natural History (USNM) by the se- 
nior author. After obtaining specimens from other col- 
lections, there was substantially more material 
available for study, especially from Mexico, than what 
Richards had seen. 


MATERIALS AND METHODS 


A total of 2319 specimens were examined. The in- 
stitutions from which specimens were examined are 
presented in Table 1. 

Material was identified by using the revisions of 
Richards (1936b, 1941) and by comparing his identi- 
fied material with the unknown specimens. Richards’ 
material included specimens of B. deducta that were 
compared with the types at The Natural History Mu- 
seum, London (BMNH), and the types of B. similaris, 
B. mexicana, B. schausi, and B. morelosa at the USNM 
were examined. 


Lepidoptera, Noctuidae, Bulia, cladistics, key to species, larval plant hosts, Euphorbiaceae, Fabaceae. 


After removing the abdomen from the specimen it 
was placed in a 10% solution of postassium hydroxide 
(KOH) and heated in a Thermolyne® Type 17600, 
Dri-Bath for approximately 20 minutes or until the 
abdomen was soft and the muscles dissolved. The 
abdomen was descaled and the internal organs re- 
moved, The specimen was then identified and the ab- 
domen, with genitalia attached, was placed in a geni- 
talia vial containing glycerin and pinned beneath the 
specimen. 

For specimens that were slide mounted, the ab- 
domen was stained in an aqueous solution of Chlorazol 
Black E. The male aedoeagus was removed from the 
genitalia and the vesica was inflated using a syringe 
filled with 99% isopropyl alcohol. The abdominal pelt 
and the rest of the genitalia were placed in solutions of 
40% and 70% ethanol for an hour. The abdomen, gen- 
italia, and aedoeagus were then stained in Eosin Y dis- 
solved in 99% isopropyl alcohol until the desired color 
was achieved. The female genitalia were treated in a 
similar manner. The genitalia and abdomen were then 
mounted on a slide using Canada Balsam. 


PHYLOGENETICS 


The taxa analyzed included 5 species of Bulia and 
two outgroup taxa, Melipotis jucunda (Hiibner) 1818 
(type species of Melipotis Hiibner 1818) and Drasteria 
fumosa (Strecker) 1898. The outgroups were selected 
based on the results of Richards (1933, 1936b), who 
included these genera and Litocala Harvey 1875 and 
Hypocala Guenée 1852 as his “group 3” and stated 
that “This is probably the best defined of all the ere- 
bine groups.” This group is based on shared characters 
of the thoracic tympanum (Richards 1933): (1) nodular 
sclerite shape, (2) pocket I pouched, and (3) pocket IV 
flanged. 


130 


deducta 


similaris 


schausi 


mexicana 


confirmans 


Drasteria 


Melipotis 


Fic. 1. Cladogram of the species of Bulia. Synapomorphies re- 
fer to solid black rectangles with character numbers on top and state 
numbers on bottom. Tree length 25, consistency index (C.I.) 1.00, 
retention index (R.I.) 1.00. 


Characters and states. All characters were run as 
ordered with their inferred plesiomorphic and apo- 
morphic states shown in Appendix 1. Plesiomorphic 
states were inferred by outgroup comparison. A total 
of 16 characters were used including 11 binary and 5 
multistate characters (Appendix 1). Data were ana- 
lyzed using the Hennig86 parsimony program written 
by Farris (1988, Version 1.5). The command “mhen- 
nig*” was used for the data matrix in Appendix 2, and 
all characters were treated as ordered. The “mhen- 
nig*” command constructs several trees, each by a 
single pass, adding the taxa in a different sequence 
each time, and then applies branch-swapping to each 
of the trees, retaining just one tree for each initial one 
(Lipscomb 1994). 

Results. Applying “mhennig” to the data matrix 
(Appendix 2) resulted in a completely resolved tree 
with a length of 25, a consistency index (C.1.) of 1.00, 
and a retention index (R.I.) of 1.00. 

The resulting cladogram is shown in Fig. | and illus- 
trates the monophyly of Bulia based on the following 
characters: (1) head with an elongate projection from 
vertex (Fig. 3); (2) eighth tergum mostly membranous, 
with a narrow X-shaped tergite (Fig. 31); (3) eighth 
sternum mostly membranous, with a wine glass shaped 
sternite (Fig. 32); (4) clasper absent from valva (Figs. 
34-38); (5) clavus absent from valva (Figs. 34-38); (6) 
coremata of male arising from base of eighth tergite 
(Fig. 33); and (7) ventral plate of ostium bursa in fe- 
male attached to seventh sternum (Figs. 49-53). 

Bulia confirmans (Walker) is the most plesiomor- 
phic species for the analyzed characters and occurs in 
the Caribbean and northern South America; the re- 
maining species of Bulia are distributed from the 
southwestern United States, throughout Mexico to 
Costa Rica. Bulia confirmans is the only species which 
can be identified without resorting to genitalia and the 
larvae feed on Euphorbiaceae, while the rest of the 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TaBLE 1. Acronyms of institutions from which specimens were 
examined. 


AMNH_ American Museum of Natural History, New York, New 
York 

BMNH The Natural History Museum, London 

BrM Bryant Mather, Clinton, Mississippi 

CNC Canadian National Collection, Ottawa, Ontario 

INBIO _ Instituto Nacional de Biodiversidad, Costa Rica 

SDNH San Diego Natural History Museum, San Diego, Cali- 


fornia 
UCB Essig Museum of Entomology, University of California, 
Berkeley, California 


UNAM Instituto de Biologia, Universidad Nacional Auténoma de 
México, México City 
USNM National Museum of Natural History, Washington, D.C. 


Bulia species whose host plants are known feed on 
Fabaceae. Bulia mexicana shares the short apex of the 
sacculus with B. confirmans (character 11, state 2). 
The elongate apex of the sacculus unites schausi, sim- 
ilaris, and deducta (character 11, state 3). Autapomor- 
phies for B. schausi include the large costal lobe of 
sacculus with a slightly bifurcate apex (character 10, 
state 2) and the vesica containing one large spine 
(character 14, state 2). The V-shaped dorsal margin of 
the juxta in the male genitalia is diagnostic for B. de- 
ducta (character 12, state 2). 


LARVAL Host PLANTS 


Three species of Bulia have been reared. A single 
female specimen of B. confirmans was reared from a 
larva that bored into the stem of Jatropha gossypiifolia 
(L.) (Euphorbiaceae) from Nueva Esparta, Isla de 
Margarita, Venezuela. Jatropha gossypiifolia occurs 
throughout Mexico, the Caribbean, and South 
America from Venezuela south to Brazil, Bolivia, and 
Paraguay. This distribution completely overlaps the 
known localities of B. confirmans. Two other Bulia 
species with known host plants feed on Prosopis 
(Mesquite) (Fabaceae). Bulia mexicana was reared on 
Prosopis juliflora (Sw.) DC. from Area de Conser- 
vacion Guanacaste, Sector Santa Rosa, Estero 
Naranjo, Guanacaste Province, Costa Rica. Prosopis 
juliflora is distributed from western and southern 
Mexico through Central America to Venezuela, 
Colombia, Ecuador, and northern Peru. This distribu- 
tion overlaps that of the known distribution of B. mex- 
icana. Bulia deducta (Morrison) was reared from an 
unknown species of Prosopis from the vicinity of Pre- 
sidio, Presidio County, Texas. There are two species, 
Prosopis glandulosa Torr. and Prosopis pubescens 
Benth., that occur in the Presidio area (Simpson 
1988). Prosopis glandulosa is the most widely distrib- 
uted of these two species extending from southern 
Kansas, west to southern California and southern 


VOLUME 56, NUMBER 3 


Rq 3 Ro 
R5 R4 


131 


Fics. 2-6. 2, Wing venation of B. confirmans. 3, Descaled head of male B. confirmans, arrow indicates elongate median projection. 4, 
Broad scales enclosed in median projection. 5, Narrow scales enclosed in median projection. 6, Mesothoracic leg with hair pencil from apex of 


mesothoracic tibia. 


Utah, south through Texas, throughout northern Mex- 
ico, including Baja California, and down the east coast 
of Mexico through Veracruz and Yucatan. The range of 
Prosopis pubescens is from southwest Texas west to 
southern California and southwest Utah, south to 
northern Mexico including Baja California, Sonora, 
Chihuahua, and Coahuila. Based on the distributions 
of these species and that of B. deducta it is likely that 
P. glandulosa is the plant host of B. deducta. The other 
species of Bulia may also feed on Prosopis. The distri- 


bution of Bulia similaris Richards overlaps that of P. 
glandulosa and is sympatric with B. deducta in south- 
ern California and southern Texas. More rearing needs 
to be done in the southwestern United States to see if 
Bulia species are restricted to Prosopis or if they are 
on any related plant species. 


Bulia Walker 


Biula Walker [1858] 1857:1169 [type species: Biula 
propira Walker by monotypy.] Preoccupied by Biula 


132 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fics. 7-8. 7, Scaled head of male B. deducta. 8, Scaled head of female B. deducta showing differences in labial palps. 


Walker [1858] 1857, Notodontidae. Nye, 1975:91; 
Poole 1989:163. 

Bulia Walker 1858:1815 Proposed as an objective re- 
placement name for Biula Walker [1858] 1857. 
Richards 1936a:431; Richards 1936b:365; McDun- 
nough 1938:132:; Richards 1941-255; Nye, 1975:91; 
Franclemont and Todd 1983:125; Poole 1989:180: 
Poole and Gentili 1996:729. 

Arsisaca Walker [1866] 1865:1261 [type species: Ar- 
sisaca bolinalis Walker by monotypy.] Nye, 1975:62 
[junior synonym of Bulia]. Poole 1989:126. 

Cirrhobolina Grote 1875:117 [type species: Syneda 
deducta Morrison by subsequent designation by 
Kirby 1875:442.| Druce 1889:358; Smith 1891:58: 
Smith 1893:325; Dyar 1902 [1903]:222: Holland 
1903:259: Barnes and McDunnough 1917:85; 
Richards 1936a:431 [junior synonym of Bulia); 
Richards 1941:255; Nye 1975:123; Poole 1989:253. 

Cirrhbolina; Dyar 1902 [1903]:222. An incorrect sub- 
sequent spelling. 


9 


Fics. 9-10. Adults of B. confirmans. 9, 3, Venezuela, Trujillo, Valera (USNM). 10, °, Dominican Republic, Los Hidalgos (USNM). 


Diagnosis. Bulia species can be confused with 
some species of Drasteria, Melipotis, and Forsebia. 
The best character to separate Bulia from these gen- 
era is the presence of an elongate median projection of 
the vertex (Fig. 3). The hindwing anal lunule is yellow 
in all North American species of Bulia and the anal 
lunule in the other genera, if present, is white. Bulia 
confirmans has a white anal lunule, but the brown bor- 
der is narrower than in the other genera (Figs. 9-10). 
Bulia confirmans is also much smaller in forewing 
length (less than 15 mm) than the most similar species 
in the other genera (greater than 15 mm). Bulia is dis- 
tinct from the other genera in the male genitalia. The 
valva lacks internal armature, such as the clasper, 
clavus, and ampulla, while these are all present in the 
outgroup genera and the coremata arises from the 
base of the eighth abdominal segment in Bulia (Fig. 
33), but in the other genera the coremata (when pres- 


ent) arises from the base of the valva. 
Description. Head: Vestiture rough, scales narrow, hairlike. 
Frons base bare, forming a triangle shaped area; frontal prominence 


VOLUME 56, NUMBER 3 


133 


Fics. 11-14. Adults of B. mexicana. 11, 3, Mexico, Jalisco, Estacion de Biologia, Chamela (UNAM), 12, 2, Costa Rica, Guanacaste, Area 
de Conservacién Guanacaste, Sector Santa Rosa, Estero Naranjo (D. H. Janzen & W. Hallwachs rearing voucher no. 97-SRNP-219). 13, 2, 
Costa Rica, Guanacaste, Area de Conservacién Guanacaste, Sector Santa Rosa, Estero Naranjo (D. H. Janzen & W. Hallwachs rearing voucher 
no. 97-SRNP-85). 14, °, Costa Rica, Guanacaste, Area de Conservacion Guanacaste, Sector Santa Rosa, Estero Naranjo (D. H. Janzen & W. 


Hallwachs rearing voucher no. 97-SRNP-91). 


present; scale tuft of frontal prominence directed dorsad; scales lat- 
eral to frontal prominence tuft directed dorsad and curved medially. 
Labial palpus appressed to front; middle segment elongate, more 
than 3 times length of basal segment; apical segment longer than 
basal segment, less than a third length of middle segment; apical 
segment wider in male than female and more concealed by scales of 


both the median and apical segments, female apical segment narrow 
and prominent. Eye well developed. Ocelli present. Vertex with a 
pair of elongate (longer than head) triangular shaped projections 
(Fig. 3) that enclose 2 types of scales (Figs. 4-5) and a median elon- 
gate spine with a slightly decurved apex; projections invaginated into 
vertex. Vertex with lateral hairlike scale tufts from near antennal 


Ve 


: 


\ 


\F 


Pe ie 18 


Fics. 15-18. Adults of B. schausi. 15, 3, Mexico, Districto Federal (UNAM). 16, 3, Mexico, Veracruz, Orizaba (USNM). 17, 2, Mexico, 
Districto Federal (UNAM). 18, °, Mexico, Districto Federal, Ajuco (UNAM). 


134 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 19-24. Adults of B. similaris. 19, 5, California, San Diego Co., Oriflamme Canyon, Anza Borego State Park (USNM). 20, d, Texas, 
Nueces Co., N. Padre Island (USNM). 21, 9, Mexico, Baja Califomia Norte, 14.4 mi S Campo Alfonsina (SDNH). 22, °, Mexico, Baja Cali- 
fornia Norte, 2.8 mi S of Catanina (SDNH). 23, °, Mexico, Baja California Norte, 1.2-5.4 mi § Santa Ines (SDNH). 24, °, Mexico, Baja Cali- 


fornia Norte, 11.5 mi SW San Miguel de Comondu (SDNH). 


scape, curved medially to partially conceal triangular shaped projec- 
tion; a median scale tuft from base of triangular shaped projection, 
extending over projection. Female vertex with flat scales arranged in 
a triangular pattern. Antenna filiform; setose ventrally. Proboscis 
well developed. Thorax: Smoothly scaled. Proleg with epiphysis 
present on tibia, less than half length of tibia. Mesotibia with elon- 
gate (more than 3/4 length of tibia) hair pencil arising from a spe- 
cialized scale patch laterad at proximal apex, hair pencil enclosed in 
a laterad elongate groove (Fig. 6); spines absent; spurs with shortest 
less than half length of longest. Mesotibia with 2 pairs of tibial spurs; 
proximal pair with shortest more than half length of longest; distal 
pair with shortest less than half length of longest; spines absent. 
Forewing: Length 12-19 mm. Vein R1 from discal cell, ending on 
costal margin; R2 and R3 + 4 from areole; R2 ending on costal mar- 
gin; R3 and R4 long stalked, ending on costal margin; R5 from are- 
ole, connate with R3 + 4, ending on outer margin; M1 from upper 
angle of discal cell; M2 and M3 from anal angle of discal cell; M3 
nearer M2 than CuA1. Hindwing: Length 10-13 mm. Vein Sc + R1 
to apex; Rs and M1 connate; M2 present well developed from anal 
angle of discal cell; M3 and CuA1 connate; 2A and 3A present. Ab- 


domen: Smooth scaled. Male with seventh tergite with a small, me- 
dian, pointed projection on proximal margin. Highth tergite with a 
narrow X-shaped tergite, remainder membranous (Fig. 31). Eighth 
sternum with a wine glass shaped sternite that forms a small triangle 
at proximal end bearing a pair of tenuis lateral arms and a median 
elongate arm extending dorsad into wide intersegmental membrane, 
remainder membranous (Fig. 32); a pair of elongate coremata from 
lateral arms, bifurcate and covered with fine hairs (Fig. 33). Female 
abdomen without modifications. Male Genitalia: Uncus well de- 
veloped, base broader than apex, lateral setae at middle. Scaphium 
as long as uncus well sclerotized. Subscaphium well developed, nar- 
rower than scaphium, striate. Valva elongate, narrow, length greater 
than 6 times width. Cucullus truncate and not as sclerotized as valva. 
Sacculus well developed, extends beyond apex of valva; costal lobe 
prominent; apex produced into a dorsally curved spine. Juxta well 
developed, excavated dorsad. Clasper, ampulla, and valvula absent. 
Aedoeagus slender, well sclerotized. Vesica well developed with 
multiple diverticula, spines, and cornuti of two types, (1) variously 
sized, spinelike and (2) elongate, hairlike. Female Genitalia: Sev- 
enth sternite indented medially at apex, with or without a median 


VOLUME 56, NUMBER 3 


135 


Fics. 25-30. Adults of B. deducta. 25, 4, Arizona, Cochise Co., Peloncillo Mts., Guadelupe Canyon (USNM). 26, 2, Arizona, Cochise Co., 
Cherry Canyon, near Cherry (USNM). 27, °, Texas, Cameron Co., San Benito (USNM). 28, 2, Texas, Brewster Co., Alpine (USNM). 29, °, 
Arizona, Cochise Co., Huachuca Mts., Ash Canyon Road (USNM). 30, 2, Texas (USNM). 


prong. Ostium at apex of seventh sternite. Highth segment tele- 
scopes inside seventh. Eighth sternite not joined medially. Segment 
9 and 10 membranous and longer than segment 8. Apophyses pos- 
teriores elongate, extending beyond proximal margin of segment 8. 
Apophyses anteriores elongate, extending to or beyond middle of 
segment 7. Papillae analis sparsely setose, apex produced. 


Discussion. The only species in Bulia that can be 
readily separated from the others is B. confirmans,with 
its small size and white anal lunule in the hindwing. All 
other species of Bulia, with the yellow anal lunule in 
the hindwing, can only be identified by genitalic char- 
acters. 

Other species with which Bulia species potentially 
can be confused are Drasteria eubapta Hampson, D. 


fumosa (Strecker), D. pallescens (Grote & Robinson), 
Forsebia perlaeta (H. Edwards), Melipotis novanda 
(Guenée), and M. indomita (Walker). 


KEY TO SPECIES 


1. Hindwing with anal lunule white (Figs. 9-10) .... confirmans 
Hindwing with anal lunule yellow (Figs. 11-30) .......... D} 
2. Head with large median projection of vertex, can be con- 
cealed by hairlike scale tufts; (Fig. 3); apical segment of labial 
palpus short and partially concealed by scales of middle 
SaqMene (MS, Tp WHMIS oo caponccoogecoanoovvcx0gcusn00® 
2’, Head without large median projection of vertex, scales flat 
not tufted; apical segment of labial palpus long and not con- 
cealed by scales of middle segment (Fig. 8); female ....... 6 
3. Juxta with dorsal margin V-shaped; lateral projections scobi- 
nate, pointed and heavily sclerotized (Figs. 38 and 43) deducta 


136 


Fics. 31-33. 


& 


OUR 


OF 


6’. 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


SO 


Structures associated with the abdomen. 31, 3, Eighth abdominal stemite of B. confurmans (genitalia slide USNM 46368). 32, 
, Eighth abdominal tergite of B. confirmans (genitalia slide USNM 46368). 33, 3, Coremata of B. similaris (genitalia slide USNM 46332). 


Juxta with dorsal margin U-shaped, lateral projections absent 


(ales, GS=O7 enavel HO) on cocdocccsccncccnonecoancens 4 
Vesica with a single large spine (Fig. 46) ...........: schausi 
Vesica with two large spines (Figs. 45 and 47) ............5 


Costal lobe of sacculus small, triangulate (Fig. 35); disto- 
lateral diverticulum elongate, more than twice length of api- 
cal spine (Fig. 45); base of vesica lacking small pair of ventral 
Giverticul ay (Hica/.o) anne nner en ert mexicana 
Costal lobe of sacculus large, truncate to triangulate (Fig. 37); 
disto-lateral diverticulum short, less than twice length of api- 
cal spine (Fig. 47); base of vesica with a pair of small ventral 
Gtivermomley (Iter, 2) oc ccoccsecccececce000s0 0000! similaris 
Seventh sternite with median prong elongate, extending above 
apex of lateral prolongation (Figs. 53 and 58) ....... deducta 
Seventh sternite with median prong short, extending to or 
below apex of lateral prolongation (Figs. 50-52 and 55-57) 7 
Corpus bursae with two sclerotized structures, (1) an elon- 
gate curved process with pointed apex at base of ductus sem- 
inalis, (2) ventral to 1 a sclerotized area with a curved apex 
pointed in the opposite direction of 1 (Fig.51) ......: schausi 
Corpus bursae lacking large sclerotized structures (Figs. 50 
and 52) 


8. Seventh sternite with median prong short, height less than 
width (Hig 00) och ons epee ce ae mexicana 

8’. Seventh sternite with median prong elongate, height greater 
thankwidthy (Ei catol/) len eet eee eee eee simiilaris 


Bulia confirmans (Walker) 
(Figs. 2-6, 9-10, 31-32, 34; 39, 44, 49, 54, 59) 

Bolina confirmans Walker [1858] 1857:1157. 

Bolina umbrosa Walker [1858] 1857:1158. [Syn- 
onymized by Richards 1936a:433.] 

Biula propira Walker [1858] 1857:1170. [Synonymized 
by Richards 1936a:433. | 

Arsisaca bolinalis Walker [1866] 1865:262, new syn- 
onym. 

Bulia bolinalis, Poole 1989:180. 

Bulia confirmans, Richards 1936a:433; Richards 
1939:pl. V, Figs. 7-9; Richards 1941:258; Poole 
1989:180. 


VOLUME 56, NUMBER 3 17 


Fics. 34-38. Male genitalia. 34, B. confirmans (genitalia slide USNM 46366). 35, B. mexicana (genitalia slide USNM 46333). 36, B. 
schausi (genitalia slide USNM 46362). 37, B. similaris (genitalia slide USNM 46323). 38, B. deducta (genitalia slide USNM 46322), cl = costal 
lobe of sacculus, sa = sacculus apex. 


Fics. 39-43. Male juxta. 39, B. confirmans (genitalia slide USNM 46366). 40, B. mexicana (genitalia slide USNM 46333). 41, B. schausi 
(genitalia slide USNM 46362). 42, B. similaris (genitalia slide USNM 46323). 43, B. deducta (genitalia slide USNM 46322). 


138 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 44-48. Male aedoeagus. 44, B. confirmans (genitalia slide USNM 46366). 45, B. mexicana (genitalia slide USNM 46333). 46, B. 
schausi (genitalia slide USNM 46362). 47, B. similaris (genitalia slide USNM 46323). 48, B. deducta (genitalia slide USNM 46322). 


Diagnosis. This is the most easily recognized 
species of Bulia. It is the only one with a white lunule 


on the hindwing, all other Bulia have a yellow lunule. 
Description. Adult male. Head: Frons white with light brown 
scales. Vertex light brown. Labial palpus with basal segment mostly 
white with some brown scales laterally; middle and apical segments 
brown with some white. Antennal scape brown, light brown later- 
ally; scaled dorsally, setose ventrally, setae approximately width of 
antennal segments. Thorax: Patagium, mesothorax, and metathorax 
light brown. Prothoracic femur brown with some white scales; tibia 
brown with an indistinct median white band, apical band white; tarsi 
brown with white apical rings. Mesothoracic femur white speckled 
with brown; tibia brown speckled with white; tarsi brown with white 
apical rings. Metathoracic femur white speckled with brown; tibia 
brown speckled with white dorsally, white ventrally, a tuft of scales 
below proximal apex buff; tarsal segment 1 brown and white, with 
white apical ring, rest brown speckled with white, white apical rings. 
Underside white. Forewing: Length 11.5-13.5 mm. Basal patch 
light brown; costal margin brown; basal band brown to M vein; me- 


dian line black to R vein; median band cream to middle of discal 
cell, speckled with brown to costal margin; orbicular spot a small 
black patch; two indistinct whitish scale patches beyond orbicular 
spot; reniform spot cream; postmedial line black from posterior 
margin, angled back toward outside of median band and bordering 
outside of reniform spot to its apex; postmedial band brown; subter- 
minal line buff from termen to M1 cell: terminal line black, scal- 
loped, from termen to M1 vein. Underside ground color white; 
brown patch at end of discal cell; brown band on terminal third. 
Hindwing: Ground color white; band on terminal fourth brown; 
white spot at apex of CuA1 cell. Underside as in upperside; white 
apical spot larger. Abdomen: Light brown dorsally; white ventrally. 
Genitalia (Figs. 34, 39, 44): Sacculus with costal lobe triangular; 
apex a short projection. Juxta with U-shaped dorsal margin. Aedoea- 
gus without dorsal spiculations near apex. Vesica with a large and a 
small spine; numerous minute cornuti. (All attempts at inflating 
vesica failed.) 

Adult female. Essentially as described for male except: 
Forewing: Length 13.0-14.5 mm. Basal patch brown speckled with 
white and light brown; median band indistinct, speckled with light 


VOLUME 56, NUMBER 3 


139 


5S 


Fics. 49-53. Female genitalia. 49, B. confirmans (genitalia slide USNM 46367). 50, B. mexicana (genitalia slide USNM 46378). 51, B. 
schausi (genitalia slide USNM 46379). 52, B. similaris (genitalia slide USNM 46325). 53, B. deducta (genitalia slide USNM 46327). 


brown, white, and brown; reniform spot speckled with light brown, 
white, and brown; subterminal line from termen to costa; terminal 
line from termen to costa. Abdomen: Underside white, pale brown 
to buff at apex. Genitalia (Figs. 49, 54): Seventh segment deeply in- 
vaginated; median prong absent. 

Type material. Bolina confirmans Walker; holotype 9°, St. 
Domingo [Dominican Republic], in BMNH. Bolina umbrosa 
Walker; holotype 2, St. Domingo [Dominican Republic], in BMNH. 


Biula propira Walker; holotype ¢, St. Domingo [Dominican Repub- 
lic], in BMNH. Arsisaca bolinalis Walker; holotype 3, Jamaica, in 
BMNH. 

Material examined. 47 ¢ and 102 °. All material is from the 
USNM unless otherwise noted. COLOMBIA: ATLANTICO: Cua- 
tro Bocas, 22 Jan. 1959, J.F.G. Clarke, 1 6. CUBA: No specific lo- 
cality, 2 2; Santiago, 1 d, 1 9. CURACAO: 3 km NE Willemstad, 
8-15 Feb. 1987, 1 2, 13 Feb. 1987, 2 2°, W.E. Steiner & JM. 


140 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 54-58. Female ventral plate of ostium bursa. 54, B. confirmans (genitalia slide USNM 46367). 55, B. mexicana (genitalia slide USNM 
46378). 56, B. schausi (genitalia slide USNM 46379). 57, B. similaris (genitalia slide USNM 46325). 58, B. deducta (genitalia slide USNM 46327). 


Swearingen; 3 km NW Fuik, 11 Feb. 1987, W.E. Steiner & J.M. 
Swearingen, | 6. DOMINICAN REPUBLIC: LA VEGA 
PROVINCE: Constanza, Hotel Nueva Suiza, 1164m, 29 May 1973, 
D. & M. Davis, 1 d, 1 2. MONTE CRISTI PROVINCE:10 km S 
Monte Cristi, 5 m, 23 May 1973, D. & M. Davis, 10 d, 36 2; Los Hi- 
dalgos, 4-5 June 1969, Flint & Gomez, 6 2; Mt. Cristy, Santo 
Domingo, Atwater, 1 °; Santo Domingo, Atwater, | ¢. GRENADA: 
Hardy Bay, 26 Oct.—4 Nov. 1975, E.L. Todd, 2 4, 3 9; Lance aux 
Epines, 30 Oct. 1975, E.L. Todd, 1 °. HAITI: No specific locality, 1 
3. JAMAICA: No specific locality, Col. B. Neumogen, 2 ¢, 1 2, Col- 
lection J.B. Smith, 1 ¢, 1 9, Collection Wm Schaus, 3 °, Collection 
Brklyn Mus, 1 9, Col. E.L. Graef, 1 J, 1 °. CLARENDON PARISH: 
2km S Rocky Pt., nr. Jackson Bay Cave, 5 m, 10 Dec. 1975, D. & M. 
Davis, 1 2, 1 2; Portland Ridge, nr. Jackson Bay Cave, 40 ft., 4 May 
1973, D. & M. Davis, 6 ¢,7°, ° genitalia slide USNM 46367; ST. AN- 
DREW PARISH: Upper Mt. View, July 1942, 2 9, July 1948, 1°, Aug. 
1942, 1 6, d genitalia slide USNM 46366, C.B. Lewis. ST. ANN 
PARISH: nr. Runaway Bay Cave, 50 tht, L=® May 1973, D. & M. 
Davis, 1 °; Hill Gardens, 14 June 1923, C.O. Gowdy, 1 6; Kingston, 
about 70 ft., 4-10 Dec. 1910, 136 (AMNH); Mandeville, Manches- 
ter, about 2250 ft., 18-20 Jan. 1920, 1 d (AMNH); Port Royal, 27 Feb. 
1911, 1 ° (AMNH). PUERTO RICO: Isla Maguey, Parguera, 20 
Dec. 1962, P. & P. Spangler, 2 2. VENEZUELA: ARAGUA: Rancho 
Grande, 1100 m, cloud forest, 30-31 Mar. 1978, J.B. Heppner, | 2. 
LARA: EI Cuji, 7 mi N Barquisimeto, 29 June-1 July 1967, R.W. 
Poole, 1 2; Puente Torres, 24 km E Carora, 10 Mar. 1978, thorn forest, 
J.B. Heppner, 9 4, 16 2, d genitalia slide USNM 46368. MERIDA: Rio 


Chama Valley, 12 km SW Ejido, 4000 ft., dry forest, 17 Feb. 1978, J.B. 
Heppner, | ¢, 7 2. ZULIA: El Tucuco, Sierra de Perijé, montane forest, 
28-29 Jan. 1978, J.B. Heppner, 2 2. Isla de Margarita, Nueva Esparta , 
23 June 1997, R. Segura, stem borer Jatropha gossypiifolia, 1 4; Valera, 
E.P. de Ballard, 1 4; Valera, Trujillo, Nov. 1922, H. Pittier, 1 6. 

Larval host. Jatropha gossypiifolia (L.) (Euphorbiaceae). This 
species is considered a noxious weed in Puerto Rico (GRIN Data- 
base 2000). The distribution of J. gossypiifolia is from Mexico south 
to Honduras and Nicaragua, throughout the Caribbean to Venezuela 
and Colombia south to Ecuador and Brazil. Bulia confirmans com- 
pletely overlays this distribution. The single specimen reared was 
from Isla de Margarita, Nueva Esparta, Venezuela. 

Distribution (Fig. 59). Collections from the Caribbean include 
localities in Cuba, Jamaica, Haiti, Dominican Republic, Puerto Rico, 
Grenada, and Curacao and in South America from northern 
Venezuela and Colombia. 


Discussion. The female of B. confirmans is more 
variable in maculation than the male. Specimens from 
the Dominican Republic and Jamaica show a pattern 
(Fig. 10), which is less distinct, but has all of the pat- 
tern elements that are present in the male. In 
Venezuela the patterned female is present, but un- 
common. The most common female there is com- 
pletely without pattern, except for a black orbicular 
spot and terminal line. The ground color is brown. 


VOLUME 56, NUMBER 3 


ae 5 QS 


~ 


wi = 4 
2a ua wh 
<i =I 44} omy 
We F; ¥ Kay 
\ Me 4 \ 


Fic. 59. Known localities from collections of B. confurmans (solid 
circles), B. mexicana (open squares), B. schausi (solid triangles). 


An adult specimen of B. confurmans was sent to the 
senior author for identification as a possible biological 
control agent for J. gossypiifolia. 


Bulia mexicana (Behr) 
(Figs. 11-14, 35, 40, 45, 50, 55, 59) 
Syneda mexicana Behr 1870:26. 


Bulia mexicana, Richards 
1941:258: Poole 1989:180. 


1936a:433; Richards 


Diagnosis. The remaining species of Bulia are almost 
indistinguishable from each other except for characters 
in the male and female genitalia. Bulia mexicana is sepa- 
rated from the other species by the following combina- 
tion of characters in the male genitalia (Fig. 35): (1) juxta 
U-shaped; (2) sacculus lobe small, triangular; (3) ae- 
doeagus with external spiculations absent; and (4) disto- 
lateral diverticulum longer than aedoeagus. The most 
closely related species to B. mexicana is B. schausi, but 
the females of B. mexicana can be differentiated from 
the females of B. schausi by the absence of any heavily 
sclerotized areas in the corpus bursae (compare Figs. 50 
and 51); and females of B. mexicana can be separated 
from females of B. similaris and B. deducta by the me- 
dian prong of the seventh segment being wider than 
high in B. mexicana (compare Figs. 50 and 57-58). 

Description. Adult male. Head: Frons white with light brown 
scales. Vertex light brown. Labial palpus with basal segment mostly 
white with some light brown scales laterally; middle and apical seg- 
ments light brown with some white. Antennal scape brown, white 
laterally; scaled dorsally, setose ventrally, setae approximately width 
of antennal segments. Thorax: Patagium, mesothorax, and metatho- 
rax brown. Prothoracic femur brown with some white scales; tibia 
brown with some white scales, apical band white; tarsi brown with 
white apical rings. Mesothoracic femur white speckled with brown; 
tibia with equal amounts of brown and white; tarsi brown with white 
apical rings. Metathoracic femur white speckled with brown; tibia 
with equal amounts of brown and white, white ventrally, a tuft of 
scales below proximal apex buff; tarsal segment 1 brown and white, 
with white apical ring, rest brown speckled with white, white apical 
rings. Underside white. Forewing: Length 14.0-16.0 mm. Basal 
patch brown; costal margin brown; basal band black to M vein; me- 
dian line black to R vein; median band cream to middle R vein, light 


141 


Fic. 60. Known localities from collections of B. similaris. 


brown to costal margin; orbicular spot a small black patch; two in- 
distinct whitish to pale gray scale patches beyond orbicular spot; 
reniform spot cream; postmedial line black from posterior margin, 
angled back toward outside of median band and bordering outside 
of reniform spot to its apex; postmedial band brown with pale gray 
median stripe; subterminal line cinnamon from termen to M1 cell; 
terminal line black, scalloped, from termen to apex. Underside 
ground color cream; wide brown patch at end of discal cell from R 
to 1A + 2A vein; brown band on terminal third. Hindwing: Ground 
color light brown; band on terminal two-fifths brown; lunule yellow. 
Underside ground color cream; small brown spot in middle of discal 
cell; brown terminal band; lunule cream. Abdomen: Light brown 
dorsally; white ventrally. Genitalia (Figs. 35, 40, 45): Sacculus with 
costal lobe triangular; apex a short projection. Juxta with U-shaped 
dorsal margin. Aedoeagus without dorsal spiculations near apex. 
Vesica with a small spine at apex of lateral diverticulum, disto-lateral 
diverticulum longer than aedoeagus with large apical spine; numer- 
ous minute comnuti on all diverticula except lateral. 

Adult female. Essentially as described for male except: 
Forewing: Length 15.0-17.0 mm. Basal patch light brown speckled 
with brown; basal band black, thin, same width as postmedial line; 
median band light brown, speckled with brown; reniform spot light 
brown speckled with brown; subterminal line faint, from termen to 
costa. Abdomen: Light brown speckled with brown. Underside 
white speckled with brown, buff at apex. Genitalia (Figs. 50, 55): 
Seventh segment deeply invaginated; median prong short, width 
greater than height, does not extend above lateral projections. Cor- 
pus bursa with basal spicules larger than rest. 


Fic. 61. Known localities from collections of B. deducta. 


Type material. Bolina mexicana Behr; neotype 3, Mexico, 
Sinaloa, Mazatlan, in USNM [designated by Richards 1936a]. 

Material examined. 20d and 26 °. All material is from the 
USNM unless otherwise noted. COSTA RICA: GUANACASTE: 
Area de Conservacién Guanacaste, Sector Santa Rosa, Estero 
Naranjo, 2 é, 12 Jan. 1997, 97-SRNP-87, 97-SRNP-92, 2 3, O7- 
SRNP-85, 97-SRNP-88, 97-SRNP-91, 97-SRNP-97.1, 4 2, 23 Jan. 
1997, 97-SRNP-219, 1 °: Playa Naranjo, Santa Rosa P.N., Mar. 1991, 
INBIOCR1000674402, INBIOCR1000674399, 2°, May 1991, 1 °, 
INBIOCR1000386796, Dec. 1990, INBIOCR1000514740, 1 9°, E. 
Alcazar; R. San Lorenzo, R.F. Cord., (Tenorio), 1050m, June 1991, 
INBIOCR1000610645, GC. Alvarado, 1 2: Santa Rosa National Park, 
1-3 June 1979, INBIOCR1002581940, D.H. Janzen, 1 ®%. 
PUNTARENAS: Est. San Miguel, Punta San Miguel, 1 d, 2 Oct. 
1997, INBIOCR1002561384, F. Alvarado, 1° (INBIO). Avangarez, 
July, 1 ¢, 3 genitalia slide USNM 46377, Schaus and Barnes Coll., 
June, 1°, 2 genitalia slide USNM 46378, Wm. Schaus Coll. MEX- 
ICO: CHIAPAS: SE Tonala, 5 June 1969, 2 3,5 °, 2 genitalia slide 
MGP # 1128, A. Mutuura (CNC). COLIMA: Cuyutlan, Jan., C.C. 
Hoffmann, 1 ¢, 1 2 (AMNH). JALISCO: Estacién de Biologia, 
Chamela, 23-24 Feb. 1981, 1 d, 24-27 May 1982, 1 d, 5/12/1980, 1 d, 
1-3 Dec. 1981, 1 ¢, A. Pescador, 1-2 May 1976, 1 6, C. Beu- 
telspacher (UNAM); Navidad, 1 Apr. 1939, A.H. & S.H Rinde, 2 on 
genitalia slide 3622, 1 ° (AMNH). NAYARIT: 5 mi E San Blas, 24-26 
Apr. 1961, Howden & Martin, 4 2°; Nuevo Vallarta, 27 Dec. 1988, 
A.D. Warren, 1 ° (CSU). OAXACA: Tehuantepec, 15-16 July 1964, 2 
3, PJ. Spangler, 11 July 1955, Vaurie, 1°, ° on genitalia slide 8760 
(AMNH). SINALOA: Mazatlan, 17-23 July 1963, P.J. Spangler, 1 ¢, 
2 2, é genitalia slide USNM 46333, 9 May 1961, 2 3, 10 May 1961, 1 
3,7, ° genitalia slide MGP # 1127, Howden & Martin (CNC), 22 
July 1954, 1 ¢, M. Casier, W. Gertsch, Bradts (AMNH). 

Larval host. Specimens were reared on Prosopis juliflora (Sw.) 
DC. (Fabaceae) from Area de Conservacion Guanacaste, Sector 
Santa Rosa, Estero Naranjo, Guanacaste Province, Costa Rica. It is 
considered a noxious weed species in Puerto Rico and Hawaii 
(GRIN Database 2000). Data for these rearings can be found on the 
Caterpillar Rearing Voucher Databases for the Area de Conserva- 
cion (ACG) in northwestern Costa Rica (Janzen 2000). 

Distribution (Fig. 59). Collections of B. mexicana are from west 
central Mexico along the Pacific coast to Chiapas and northwestern 
Costa Rica. 

Discussion. The overall shade of the color pattern 
can vary between light and dark. This also occurs in 
specimens of B. similaris from Mexico. There are two 
forms in the female forewing color pattern. The macu- 
late form shows all of the color pattern elements 
that are present in the male, but not as distinct, giving 
these specimens a washed out appearance. The im- 
maculate form is almost completely lacking a pattern, 
except for the faint subterminal and terminal lines, the 
small black subapical spot and the large dark reniform 
spot. 


Bulia schausi Richards 
(Figs. 15-18, 36, 41, 46, 51, 56, 59) 


Bulia schausi Richards 1941:259: Poole 1989:180. 


Diagnosis. Bulia schausi tends to be grayer in 
ground color than B. mexicana, but dissection is nec- 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


essary for accurate identification. The differences in 
the male genitalia of B. schausi from other species of 
Bulia are the costal lobe of the sacculus is slightly bi- 
furcate (Fig. 36) and the vesica bears a single spine 
(Fig. 46). In the female genitalia, these differences are 
the corpus bursa has a heavily sclerotized cone within 
the ductus seminalis and ventral to this is a heavily 
sclerotized area that ends in a blunt apex (Fig. 51). 

Description. Adult male. Forewing: Length 15.0-16.5 mm. 
Genitalia (Figs. 36, 41, 46): Sacculus with costal lobe slightly bifur- 
cate; apex an elongate projection. Juxta with U-shaped dorsal mar- 
gin. Aedoeagus without dorsal spiculations near apex. Vesica with 
lateral diverticulum absent, disto-lateral diverticulum with large api- 
cal spine and minute comuti at base of disto-lateral diverticulum, 
cornuti larger on ventro-proximo lateral diverticula. 

Adult female. Forewing: Length 15.5-16.0 mm. Genitalia 
(Figs. 51, 56): Seventh segment deeply invaginated; median prong 
short, width greater than height, extends to lateral projections. Cor- 
pus bursa with heavily sclerotized cone within the ductus seminalis; 
ventral to this is a heavily sclerotized area ending in a blunt apex; a 
band of elongate spicules between these two areas. 

Type material. Bolina schausi Richards; holotype 2, Mexico, 
Tehuacan, in USNM. 

Material examined. 12d and 10 2. All material is from the USNM 
unless otherwise noted. MEXICO: DISTRICTO FEDERAL: 
Ajucco, 3000m, 1 June 1981, 1 °, C. Beutelspacher (UNAM); Cha- 
pultepec, 20 June 1939, 1 2°, C. Beutelspacher (UNAM); Jardin 
Botanico, C.U., 5 Nov. 1969, C. Beutelspacher (UNAM); Xochim- 
ilco, 3 July 1983, 1 d, C. Beutelspacher (UNAM); 18 June 1963, 1 d, 
21 June 1963, 1 2, 12 July 1960, 1 ¢, R. Johnson (UNAM). DU- 
RANGO: 10 mi W Durango, 15 June 1964, W.C. McGuffin, (1 2) 
(CNC); 7 mi W Durango, 26 July 1964, 1 d, ¢ genitalia slide MGP # 
1132, W.C. McGuffin (CNC); El Salto, Rcho. Nuevo, 10-13 June 
1989, Broomfield, 1 ¢ (SDNH); Vte. Gro., 3 June 1984, 1 d, 1 2 
(UNAM). MEXICO: 7 air mi WSW Juchitepec, 275m, 24 Aug. 
987, J. Brown & J. Powell, 1 ° (UCB). PUEBLA: Tehuacan, Sep. 
1908, Muller, 1 2, paratype, ° genitalia slide USNM 46379; Sep. 
1937, ° on genitalia slide 3612, 20 May 1941, d on genitalia slide 
3501, 1 3, 1 2 (AMNH). SONORA: 16 mi NE Cd. Obregon, 13-17 
May 1961, Howden & Martin, 1 4, d genitalia slide MGP# 1131 
CNC). VERACRUZ: Fortin, 7 June 1988, T. Taylor, 1 6 (CSU); 
Orizaba, 2 d, d genitalia slide USNM 46380, 2°, ° genitalia slides 
USNM 46381, 46382. 

Larval host. Unknown. 


Distribution (Fig. 59). Collections from northwest to east cen- 
tral Mexico. Localities are near or in the Sierra Madre Occidental. 

Discussion. This is the rarest species of Bulia. The 
sexual dimorphism in the female is the immaculate 
form in which there is virtually no pattern and the 
reniform spot is large and dark. 

Bulia schausi is associated with the mountainous 
central region of Mexico. It is sympatric with B. simi- 
laris in the vicinity of Tehuacan, Puebla, Mexico. 


Bulia similaris Richards 
(Figs. 19-24, 33, 37, 42, 47, 52, 57, 60) 
Bulia  similaris Richards 1936a:433; Richards 
1941:262; Poole 1989:180: Poole and Gentili 
1996:729. 


VOLUME 56, NUMBER 3 


Bulia similaris race californica Richards 1939:70; 
1941:262. [Synonymized by Poole 1989:80. ] 
Bulia morelosa Richards 1941:261: Poole 1989:180, 


new synonym. 


Diagnosis. Bulia similaris is most closely related to 
B. deducta, but can be separated by the shape in the 
juxta in the male genitalia. The juxta in B. similaris has 
a U-shaped dorsal margin (Fig. 42), whereas the juxta 
in B. deducta has a V-shaped dorsal margin (Fig. 43). 
The elongate apex of the sacculus is round in B. simi- 
laris (Fig. 37), but pointed in B. deducta (Fig. 38). The 
spiculations near the apex of the aedoeagus are larger 
and cover a smaller area in B. similaris (Fig. 47), in B. 
deducta the spiculations are smaller and cover a larger 
area (Fig. 48). The cornuti on the disto-lateral divertic- 
ulum are smaller and greater in number in B. similaris 
(Fig. 47) than in B. deducta (Fig. 48). The females can 
be separated by the shape of the median prong of the 
seventh abdominal segment. The prong width is 
greater than its height in B. mexicana and B. schausi 
(Figs. 55-56), but in B. similaris and B. decucta the 
height is greater than the width (Figs. 57-58). In B. 
similaris the length of the median prong is less than or 
equal to the height of the lateral projections (Fig. 57); 
in B. deducta the median prong is greater than the 
height of the lateral projections (Fig. 58). 


Description. Adult male. Forewing: Length 13.0-16.5 mm. 
Genitalia (Figs. 37, 42, 47): Sacculus with costal lobe triangular 
with a rounded apex; apex an elongate projection. Juxta with U- 
shaped dorsal margin. Aedoeagus with dorsal spiculations near apex. 
Vesica with largest spine on lateral diverticulum, smallest spine on 
disto-lateral diverticulum; minute cornuti ventral and lateral on 
disto-lateral diverticulum and ventro-lateral diverticula; ventro-prox- 
imal diverticula absent; largest cornuti dorsally on disto-lateral di- 
verticulum; an extra pair of small ventral diverticula at base of vesica. 

Adult female. Forewing: Length 12.5-18.0 mm. Genitalia 
(Figs. 52, 57): Seventh segment deeply invaginated; median prong 
width less than or equal to height, extending to or below lateral pro- 
jections, apex truncate or with slight V-shaped notch. Corpus bursa 
with dorso-medial invagination lightly sclerotized and containing 
fine spicules; a band of elongate spicules below junction with ductus 
bursae. 

Type material. Bolina similaris Richards; holotype 4, U.S.A., 
Texas, San Benito, in USNM. 

Material examined. 203 d and 244 °. All material is from USNM 
unless otherwise noted. MEXICO: BAJA CALIFORNIA [label data]: 
Las Parras, W.M. Mann, | d; Punta Prieta, 27 Mar. 1935, V.H. dos Pas- 
sos, 1 2; 23 mi S San Ignacio, 10 Oct. 1967, G.A. Marsh, 1 2 (UCB); 
Desengano, 9 Oct. 1962, F.T. Thorue, | 2 (UCB); L. Cantilla Cyn., 
Sierra Juarez, 20 Mar. 1967, P.A. Opler & J. Powell, 2 9 (UCB); San 
Felipe, 26 Mar. 1963, G.J. Stage, 26 (UCB); Canyon del Tajo, 1 Apr. 
1953, J. Powell, 34, 6 2 (UCB); 17 mi N Punta Prieta, 3 Oct. 1961, 
C.F. Harbison, 1 d (SDNH); 20 mi N Punta Prieta, 27 Apr. 1962, C.F. 
Harbison, 1 6 (SDNH); 8 mi N El Refugio Magdalena, 19 Oct. 1961, 
FF, Gander, 1 ¢, 2 ° (SDNH); Agua Refugio, 1 Apr. 1935, C.F. Har- 
bison, 1 2 (SDNH); Cautiles (Tajo Canyon), 20 Apr. 1955, C.F. Harbi- 
son, 1 2 (SDNH); Mesquiutal, Apr. 1928, 1 d (SDNH); Santiago, 


143 


1952, C.F. Harbison, 1 ¢d (SDNH): 15 Nov. 1936, F. Gander, 1 d 
(SDNH). BAJA CALIFORNIA NORTE: 14.4 mi S Campo Alfosina, 
20-26 Oct. 1987, N. Bloomfield, 1 ¢, 4 genitalia slide USNM 46364; 
14.4 mi S Campo Alfosina, 20-26 Oct. 1987, N. Bloomfield, 1 ¢, ¢ 
genitalia slide USNM 46364; 59 mi S Puertecitos, 14-15 Mar. 1988, 
N. Bloomfield, 1 4, 3 genitalia slide USNM 46363; Rancho Santa Ines, 
540 m, 30 June 1979, W.H. Clark, 2d, 3 2; 10 mi SE El Rosario, 31 
Mar. 1976, J. Doyen, P. Rude, R. Morrison, | 4, 2 9 (UCB); 24 mi N 
Punta Prieta, 1-2 Apr. 1973, S.L. Szerlip, J. Doyen, J.A. Powell, 1 2, 1 
Apr. 1973, 1 2, S.L. Szerlip, 2 Apr. 1973, 1 2, J.A. Powell (UCB); 7 mi 
SW Mission San Borja, 30 Mar. 1973, J.A. Powell, 1 29 (UCB); 9 km 
NW Rancho Santa Ines, 550m, 1 July 1979, W.H. Clark, 1 6, 1 2 
(CNC); Arroyo Catavina, 35 mi S El Progresso, 2 Apr. 1976, Doyen & 
Rude, 2 2 (UCB); Bahia los Angeles, 26 Dec. 1978, R.E. Dietz, 1d 
(UCB); Diablito Cyn., E face Sierra San Pedro Martir, 5-6 Apr 1973, 
J.A. Powell, 1d (UCB); 1.2 mi E Santa Ines (arroyo), 23 Mar. 1986, 
Faulkner & Broomfield, 2 2 (SDNH); 1.2-5.4 mi S Santa Ines, 5-9 
Dec. 1987, N. Broomfield, 33, 5 9 (SDNH); 1.5 mi SW San Miguel, 
16-20 Dec. 1987, N. Broomfield, 43, 6 2 (SDNH); 10.2 mi W Ranch 
Progreso (S. Francisquito), 6-7 Nov. 1987, N. Broomfield, 2 d 
(SDNH); 11.5 mi SW San Miguel de Comondu, 15 Dec. 1987, N. 
Broomfield, 3 6 (SDNH); 12 mi N Catavina, 30 Apr. 1962, C.F. Har- 
bison, 1 2 (SDNH); 13.3 mi S B. de los Angeles, 1-13 Mar. 1988, N. 
Broomfield, 4 2 (SDNH); 14.4 mi S Campo Alfonsina, 20-26 Oct. 
1987, N. Broomfield, 20 ¢, 31 9 (SDNH); 19 mi SW Campo Alfonsina 
(Canyon), 27—28 Oct. 1987, N. Broomfield, 1d, 1 2 (SDNH); 2.8 mi S 
Catavina, 23 Mar. 1981, Faulkner & Brown, 2 4, 42 (SDNH); 25.5 mi 
NE El Arco, 11-15 Dec. 1987, N. Broomfield, 1d, 1 9 (SDNH); 3 mi 
S B. de los Angeles, 2 Nov. 1987, N. Broomfield, 1 9 (SDNH); 51 mi 
S Catavina, 7 Apr. 1982, Faulkner & Brown, 1 2 (SDNH); 7 mi S Cha- 
pala, 29 Oct—1 Noy. 1987, N. Broomfield, 2 5,72 (SDNH); 9 mi E EJ 
Bonfil, 11 June 1986, Broomfield, 1 d (SDNH); 9 mi S Rosarito, 5 
Oct. 1983, Faulkner & Andrews, 1 2 (SDNH); Bahia de Los Angeles, 
30 Mar. 1983, C. Brey, 1 6 (SDNH); El Marmol, 15 Mar. 1947, 1 9, 
Harbison, 12 Apr. 1940, 2 6 (SDNH); Sierra San Pedro Martir, Las 
Encinas, 15 June 1980, Brown & Faulkner, 1 2° (SDNH); Wash, 2 mi 
N Catavina just east of Hwy 1, 3 Apr. 1992, J. Brown, 1 d, 1° (SDNH). 
BAJA CALIFORNIA SUR: 16.3 mi NE Arco, 8-10 Mar. 1988, N. 
Bloomfield, 1 4, 3 genitalia slide USNM 46365; 2 mi NW El Triunfo, 
12 Aug. 1966, J.A. Chemsak (UCB); 2 mi SE San Bartolo, 900, 9 Nov. 
1993, Y.F. Hsu (UCB); 26 mi W. La Paz, 11 Aug. 1966, J. A. Chemsak, 
1 2(UCB): 7 mi S San Pedro, 10 Aug. 1966, J. Powell (UCB); 7 mi SE 
Guerrero Negro, 8 Apr. 1976, J. Doyen & P. Rude, 2 3, 10 2 (UCB): 90 
km NW La Paz, 8 Nov. 1993, Y.F. Hsu, 1 d, 2 9 (UCB); El Coyote, 12 
Apr. 1972, R.W. Holland, 16 (AMNH); Sierra Laguna, 17 air mi ENE 
Todos Santos, 6000’, 12-18 Dec. 1979, 3 d, J. Doyen & W. Tschinkel, 
6 6, 1 2°, P Rude (UCB); 16.3 mi NE El Arco, 8-10 Mar. 1988, N. 
Broomfield, 2 3,3 2 (SDNH); 2 mi W Catavina, 4 Apr. 1935, C.F. Har- 
bison, 3 5, 4 ° (SDNH); 3 mi NE San Isidro (La Purisima), 14 Apr. 
1985, Broomfield & Faulkner, 2° (SDNH); 3.3 mi S El Cien, 26 Sep. 
1981, D. Faulkner & F. Andrews, 1 6, 2° (SDNH):5 mi N San Isidro, 
18 July 1986, Broomfield, 1 2 (SDNH); 7 mi SW Loreto, 12 Oct. 1981, 
D. Faulkner & F. Andrews, 1 2 (SDNH); 9.2 mi SSE Mulege, 23 Sep. 
1981, F. Andrews & D. Faulkner, 1 2 (SDNH); I. San Jose, NW end, 
11 July 1985, D.K. Faulkner, 1 ¢ (SDNH); La Paz, Econohotel 
Palmira, 29 Nov. 1980, Brown & Brown, 1 6 (SDNH): San Isidro, 29 
May 1984, Broomfield & Faulkner, 8 d, 3 2 (SDNH). CHIAPAS: 8 mi 
NW San Cristobal, 28 May 1969, A. Mutuura, 1¢ (CNC). 
COAHUILA: 10 mi N Monclova, 1500, 7 July 1963, Duckworth & 
Davis, 5d, 1 °, 6 genitalia slide USNM 46330; Saltillo, 7 Aug. 1963, 
PJ. Spangler, 1 2; Saltillo, 23 May 1952, d genitalia slide on 8699, 2 
genitalia slide on 8756, M. Casier, W. Gertsch, R. Schrammei, 1 d, 1 2 
(AMNH); 25 mi S Cd. Acuna, 30 June 1983, Wolfe & Valverde, 1 d, 3 


144 


2 (SDNH). CUNEO R IO: San Miguel de Allende, 14 May 1979, 
J.R. Powers, 2 ° (UCB). MORELOS: plete, 4 Feb. 1949, J.J. McK- 
elvey, 1 °. NUEVA LEON: 2 mi S Monterrey, Chipinque Mesa, 4200, 
10 Aug. 1963, Duckworth & Davis, 1 2; 3 mi E Galeana, 5000, 7-9 
Aug. 1963, Duckworth & Davis, 3 d, 2 2; Laredo, Km 1086, 375m, 29 
Noy. 1950, C. Dowling, 1 d; Monterrey, 8 Aug. 1963, P.J. Spangler, 4 d 
, 9°, d genitalia slide USNM 46331, 46332; Monterrey, 21 Mar. 1999, 
B. Mather, 1 2 (BM); 2 mi S. Monterrey, Chinpinque Mesa, 4200, 10 
pe 1963, Duckworth & Davis, 1 2 (CNC); 5 mi S. Monterrey, 10 
July 1963, 1 3, 11 July 1963, 1 4, 7 July 1963, 1 2, 13 July 1963, 1 9°, 27 
July 1963, 3°, ° genitalia slides MGP # 1124,1125, H. & A. Howden 
(CNC); Chinpinque Mesa, nr. Monterrey, 5400, 22 July 1963, 19, ° 
genitalia slide MGP # 1126, H. & A. Howden (CNC); Linares, 13 Apr. 
1954, D.H. Janzen, 1 dS (UCB): Monterrey, 9 May 1976, J.R. Powers, 
5d, 2° (UCB). PUEBLA: 2 mi SW Tehuacan, 5300, 4 Oct. 1975, 3d, 
12,5 Oct. 1975, 1d, 2 2, J.A. Powell (UCB); 7.5 km NE Azumbilla, 
2200m, 21 Aug. 1987, Brown & Powell, 1 9 (UCB). QUERETARO: 
Km 4 Carretera La Lagunita-Tilaco, 1160 m, 11 Jan. 1998, Vences & 
Ibarra, 1 2? (UNAM). SAN LUIS POTOSI: 15 mi N Valles, 18 May 
1952, ° genitalia slide on 8758 M. Casier, W. Gertsch, R. Schrammei, 
2 (AMNH); Tamazunchale, 20 May 1952, d genitalia slide on 8713 
M. Casier, W. Gertsch, R. Schrammei, 1 6 (AMNH); Xilitla, Vencidor 
Station, 22 Apr. 1977, W.H. Howe, 2 6 (AMNH). SONORA: 22 mi N 
Cd. Obregon, 11 June 1962, D.H. Janzen, 1 ° (UCB); 5 mi S Presa 
Obregon, 23 Mar. 1980, J.T. Doyen, 1 9 (UCB); Rio Yaqui, 12 mi E 
Esperanza, 300, 25 Mar. 1980, J. Doyen (UCB); 20 mi S Sonoyta, 22 
Sep. 1967, C.F. Harbison, 1 2 (SDNH); 4.3 mi E Moctezuma, 2-4 
Sep. 1987, N. Broomfield, 2 9 (SDNH). TAMAULIPAS: Guemez, 28 
June 1965, PJ. Spangler, 1 2; 4 mi SW C. Victoria, 10 July 1963, Duck- 
se & Davis, 1 ° (CNC); Gomes Farias, 3 Oct. 1986, L. Cervantes, 
2° (UNAM): Victoria, 12 Mar. 1953, 1 9, 14 Mar. 1953, 14 Mar. 1953, 

; 3, 1°, D.H. Janzen (UCB). VERACRUZ: 36 mi S Tantoyuca, 120m, 
3 July 1983, K. Wolfe & N. Valverde, 3 2 (SDNH). U.S.A.: ARI- 
ZONA: Pima Co.: Baboquivera Mts., 5000 ft., 15-30 June 1923, 1d, 1 
2, d genitalia slide made VI-3-35 A.G. Richards, 1-7 July 1923, 1 8, 
O.C. Poling. CALIFORNIA: Imperial Co.: Carrizo, 10 Mar. 1928, 
C.C. Searl, 1 6 (SDNH). Orange Co.: Silverado Canyon, Santa Ana 
Mts., 1650, 25 Apr. 1979, G.A. Marsh, 1 d (UCB). Riverside Co.: Palm 
Springs, 16-23 Apr., 14,392, ° genitalia slide USNM 40520, 29 Mar. 
1952, 1 2, 19 May 1950, 1 9°, A.H. Rindge (AMNH): Boyd Desert Re- 
search Center, 4 mi S Palm Desert, 6 Apr. 1963, J. Powell, 1 ¢ (UCB); 
Chino Canyon, Palm Springs, 19 Apr. 1950, E.C. Johnston, 1 ¢ 
(CNC); Chino Cyn, nr. Palm Springs , 20 Apr. 1960, J. Powell, 5,3 9 
(UCB). San Bemardino Co.: Joshua Tree Nat. Mon., Indian Cove, 
25-26 Mar. 1981, S.E. Miller & P.M. Mercer, 3 6; Loma Linda, 8-15 
Apr. 2d, 4°, d genitalia slide USNM 446384, ° genitalia made VI-11- 
35 A. CG. ae ae 16-23 Apr., 3d, 29, ° genitalia slide USNM 46783, 
16-23 May, 2¢, 1-7 July, 1 2; San Bernardino, 27 May 1920, E. Piazza, 
1 d; 29 Palms, 20 Apr. 1950, E.C. Johnston 1 d (CNC); Afton Rd., 23 
mi SW Baker, 23 Apr. 1977, Cave & Chemsak, 1 9 (UCB). San Diego 
Co.: Jacumba, 1, 28 May 1924, 24, 20, 24 June 1924,34,39, ° geni- 
talia slide USNM 46325, H.G. Dyar, 9 May 1924, 1 3 (CNC); Ori- 
flamme Cyn., Anza Borego State Park, 18 Apr. 1998, N. Broomfield, 9 
3,4 2, d genitalia slide USNM 46323; San Diego, 24 May 1924, 1 d, 23 
July 1924, 1 9, > al G. Dyar, 29 May 1920, 1 °, 30 June 1920, 1 d, 17 
June 1920, 1d, E. Piazza, Apr. 1°, 2 genitalia slide made VI-12-35 
A.G. Richards, Apr., 1 4, d genitalia made VI-2-35, Coquillet; Borego, 
1 May 1941, 1 6, G.H. & J.L. Sperry, Apr. 1941, 19, June 1946 12, N. 
Crickner (AMNH); La Jolla, 25 June 1963, J. Powell, 1 ° (UCB); 1 mi 
W Jacumba, 25 May 1981, C. Bruyea, 1 ° (SDNH); 2 mi E Jacumba, 
17 Aug. 1979, Brown & Faulkner, 1 d (SDNH); 5 mi N Escondito, 
250 m, 24 Apr. 1983, 1 2, 16 June 1983, 1 2, K. Wolfe & M. Valverde 
(SDNH): Boulevard-Manzanita, 3 June 1980, R. Messner, 1 6 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


(SDNH); Crouch Meadow Spr., 22 May 1979, D.K. Faulkner, 1 2 
(SDNH); Green Vly. TT Poway, 5 Apr. 1981, C. Bruyea, 1d (SDNH); 
Jacumba, 20 June 1978, Faulkner & Brown, 1 d (SDNH); Kitchen 
Creek Cyn., 25 May 1981, C. Bruyea, 1 6 (SDNH); NAS Miramar, 
Lep. Sur. Site 7, 3.5 mi E I-15,0.25 mi S Green Farms Road, 22 Apr. 
1996, N. Broomfield, 1 2 (SDNH): Rancho Bernardo Industrial Park, 
30 Apr. 1981, C. Bruyea, 1 ° (SDNH); San Diego Wild Animal Park, 
San Pasqual, 19 June 1982, G.P. Bruyea, 1 6 (SDNH). Stanislaus Co.: 
Del Rio, 26 Apr. 1959, M.R. MacKay, 2 6 (CNC). La Puerta Valley, 
July 1911, GH. Field, 2 6, 1 9 (SDNH); Vallecito Vy., 4-29, C.F. Har- 
bison, 1 ° (SDNH). TEXAS: Bexar Co.: San Aaionto. 24-30 Mar., 2d, 
2 2, d genitalia slide USNM 46324, 1-7 Apr., 1 d, 1 2, H. Schwarz. 
Brewster Co.: K- Bar Ranch, Chisos Mountains, 3400, 5 June 1973, 
D.C. Ferguson, 1 ¢; Alpine, 22 May 1950, E.C. eligi (CNC). 
Cameron Co.: San Benito, 24-30 Apr., 1 d, 8-15 May, 1d, 1 9, ° gen- 
italia slide USNM 46393, 16-23 May, 4 2. Culberson Co.: Sierra Dia- 
blo, 20 mi NNW Van Horn, 29 May 1973, D.C. Ferguson, | d. Goliad 
Co.: Goliad, Nov. 1895, 1 2, ° genitalia slide USNM 46392. Kerr Co.: 

Kerrville, 31 May 1906, 2d, d conta slide USNM 46394, F.C. Pratt. 
La Salle Co.: Artesia Wells, 11 July 1972, A. & M.E. Blanchard, 1 9; 
Cotulla, 12 May 1907, Crawford & Pratt, 2 6. Nueces Co.: Corpus 
Christi, 1 9; N. Padre Island, 9 Sep. 1974, A. & M.E. Blanchard, 2 d. 
San Patricio Co.: Sinton Welder Wildlife Foundation (San Patricio), 
22, 24-26 May 1962, 9 6, 8 9, 5 d and 2 ° genitalia slides USNM 
46385-46390, 46395, A. &M.E. Blanchard. Uvalde Co.: Garner State 
Park, 21 Sep. 1965, 1 2, 2 genitalia slide USNM 46391, A. & M.E. 
Blanchard. Victoria Co.: Victoria, 30 Mar., 1 2, 4 Apr. 1 2, E.A. 
Schwarz. Williamson Co.: eo 21 Apr. 1937, L.J. Milne, 1d 
(CNC). No specific locality,1 d, 2 °. 

Larval host. Unknown. 

Distribution (Fig. 60). Collections indicate a disjunct distribu- 
tion for B. similaris. The western distribution is from southern Cal- 
ifornia throughout Baja California, Mexico, with an isolated record 
from the Baboquivari Mountains in south central Arizona and a few 
scattered localities in northwestern Sonora, Mexico. The eastern dis- 
tribution extends from central Texas along eastern Mexico to central 
Chiapas. 

Discussion. The females of B. similaris show four 
distinct phenotypes. There are two maculate forms, a 
light one (Fig. 21), and a dark one (Fig. 22) that are 
very similar to the males (Figs. 19-20). There is an im- 
maculate form that lacks most of the pattern and has a 
large dark reniform spot (Fig. 23). A fourth form is in- 
termediate in maculation with a distinct median band 
and median line with the terminal half of the forewing 
a cinnamon ground color (Fig. 24). Richards (1939) 
described the race californica for a pale form of B. 
similaris from southem California and northern Baja 
California. In this form the overall coloration is lighter 
and the median band and reniform spot are pure 
white. The hindwing is pure white in the californica 
race, but is generally darker in the Texas and Mexican 
specimens. 


Bulia deducta (Morrison) 
(Figs. 7-8, 25-30, 38, 43, 48, 53, 58, 61) 
Syneda deducta Morrison 1875:220. 


Syneda pavitensis Morrison 1875:221. [Synonymized 
by Richards 1936:434. | 


VOLUME 56, NUMBER 3 


Cirrhobolina incandescens Grote 1875:117.[Syn- 
onymized by Richards 1936:434. | 

Cirrhobolina mexicana var. vulpina H. Edwards 
1882:14. [Synonymized by Richards 1936a:435. | 

Syneda mexicana var. albina Strecker 1900:35. [Syn- 
onymized by Richards 1936a:4:35. | 

Bulia deducta, Richards 1936a:434; Richards 
1941:263: Poole 1989:180; Poole and Gentili 
1996:729. 


Diagnosis. Bulia deducta is most likely to be con- 
fused with B. similaris. The only way to accurately 
identify these species is by examining the characters of 
the male and female genitalia. In the male, B. deducta 
can be separated from B. similaris by the shape of the 
juxta. The dorsal margin of the juxta is V-shaped in B. 
deducta (Fig. 43) and U-shaped in B. similaris (Fig. 
42). At the apex of the aedoeagus the patch of spicula- 
tions near the apex are smaller and cover a larger area 
in B. deducta (Fig. 48), and these spiculations are 
larger and cover a smaller area in B. similaris (Fig. 47). 
The cornuti on the disto-lateral diverticulum in B. de- 
ducta are larger and fewer in number (Fig. 53), and in 
B. smilaris they are smaller and greater in number 
(Fig. 52). The females can be separated by the shape 
of the median prong of the seventh abdominal seg- 
ment. In B. deducta the median prong is greater than 
the height of the lateral projections (Fig. 58), and in B. 
similaris the length of the median prong is less than or 
equal to the height of the lateral projections (Fig. 57). 

Description. Adult male. Forewing: Length 13.0-17.0 mm. 
Genitalia (Figs. 38, 43, 48): Sacculus with costal lobe large, apex 
truncate; apex an elongate projection. Juxta with V-shaped dorsal 
margin. Aedoeagus without dorsal spiculations near apex. Vesica 
with largest spine on disto-lateral diverticulum, smaller spine on lat- 
eral diverticulum; minute cornuti on apex of disto-lateral diverticu- 
lum and on ventral diverticula; patch of large cornuti medially on 
disto-lateral diverticulum. 

Adult female. Forewing: Length 12.5-18.5 mm. Genitalia 
(Figs. 53, 58): Seventh segment deeply invaginated; median prong 
width less than or equal to height, extending above lateral projec- 
tions, apex truncate. Corpus bursa with a band of elongate spicules 
below junction with ductus bursae. 

Type material. Syneda deducta Morrison; holotype ¢, U.S.A., 
Texas, Waco, in the Museum of Comparative Zoology, Harvard Uni- 
versity, Cambridge, Massachusetts. Syneda pavitensis Morrison; 
holotype 2, U.S.A., Texas; Peab. Acad.; Syneda pavitensis Type Morr. 
[Hand written in black ink], in USNM. Cirrhobolina incandescens 
Grote; holotype 2°, U.S.A., Texas, in BMNH. Cirrhobolina mexicana 
var. vulpina H. Edwards; holotype 2, U.S.A., [no locality label]; Cir- 
rhobolina mexicana Behr var. vulpina Hy. Ed.; Col. B. Neumogen; 
Collection Brklyn Mus; Type No. 34118 U.S.N.M. [Red label black 
type]; Genitalia Slide USNM 40256 [green label], 2 gen. 978, 24 
May 34 FHB [hand written black ink]; in USNM. Syneda mexicana 
var. albina Strecker; 2 syntypes °, Mexico, Chihuahua, in Field Mu- 
seum of Natural History, Chicago, Illinois. 

Material examined. 839 6 and 863 2. All material is from the 
USNM unless otherwise noted. CANADA: MANITOBA: 


145 


Cartwright, 1 6. MEXICO: BAJA CALIFORNIA [label data]: 9 mi 
SE Catavina, 16 Oct. 1967, G.A. Marsh, 1 ¢ (UCB); Gaskill’s, 14 
Nov. 1936, C.F. Harbison, 1 ¢ (SDNH). BAJA CALIFORNIA 
NORTE: Laguna Salada, 21 Sep. 1967, G.A. Marsh, 1 6 (UCB); 14.4 
mi S Campo Alfonsina, 20-26 Oct. 1987, N. Broomfield, 1 2 
(SDNH); 8 mi E Tecate, 6 July 1984, Brown and Tocco, 1 ° 
(SDNH); Cantil, Tajo Canyon, 25 Oct. 1955, Harbison, 3 d, 5 ° 
(SDNH); Tajo Canyon, 25 Oct. 1955, Harbison, 3 d, 4 2 (SDNH). 
CHIAPAS: 20 mi NE Huixtla, 900 m, 9 July 1983, Wolfe & 
Valverde, 1 29 (SDNH). CHIHUAHUA: 12 mi N Escalon, 18 Apr. 
1961, Howden & Martin, I ¢, 3 2 (CNC); 25 mi W Del Perral, 6800, 
14 July 1964, J.A. Chemsak, 1 ¢ (UCB); 42 mi N Chihuahua in cyn. 
3 mi W Parrita, 2 June 1981, J. Doyen, 1 d, 2 2 (UCB); Majalca Rd., 
25 mi NW Chihuahua, 14-17 Apr. 1961, Howden & Martin, 2 6 
(CNC); Mesa del Huracan, 7400, 21-25 July 1964, J.E.H. Martin, 2 
2 (CNC). COAHUILA: sand dunes at Bilbao, 8 mi N Viesca, 30-31 
May 1981, J. Doyen, J. Liebherr, 1 2 (UCB); 25 mi S Cd. Acuna, 300 
m, 30 June 1983, Wolfe & Valverde, 1 ¢, 3 2 (SDNH). DURANGO: 
Lerdo, 1 2; 10 mi W El Salto, 26 June 1964, 1 J, 12 July 1964, 1 2, 
W.C. McGuffin, 26 June 1964, 1 4,5 July 1964, 14, 8 July 1964, 1 9, 
J-E.H. Martin (CNC); 16 mi SE Durango, 14 July 1964, W.C. 
McGuffin, 2 2 (CNC); 5 mi W Durango, 11 June 1964, J.E.H. Mar- 
tin, 1 ¢6 (CNC); Durango, 3 May 1961, Howden & Martin, 1 ? 
(CNC); Vte. Gro., 3 June 1984, 1 2? (UNAM). NUEVA LEON: Mon- 
terrey, 25 Nov. 1909, F.C. Bishopp, 1 d, 3 genitalia made V-12-40 
A.G. Richards; Gomez Palacio, 11 May 1918, U.C. Loftin, 1 4; Chip- 
inque Mesa, nr. Monterrey, 5400’ , 22 July 1963, 12, H. & A. How- 
den, 1760’, 27-28 Apr. 1969, J.E.H. Martin, 2 2 (CNC); Linares, 9 
May 1969, 7 4, d genitalia slides MGP # 1129, 1130, 2 2° (CNC). 
SONORA: Nogales, 2 Aug. 1919, FJ. Dyar, 1 6; 95 mi E San Luis 
R.C., Hwy. 2, 4 Apr. 1966, M. Lundgren, 1 6 (UCB); Alamos,1 July 
1971, G. Kageyama, 1 6 (UCB); 4.3 mi E Moctezuma, 18-21 July 
1987, 1 °, 2-4 Sep. 1987, 3 3, 3 2, N. Broomfield (SDNH). Cuya- 
maca, 20 June 1940, 1 ° (SDNH). U.S.A.: ARIZONA: Apache Co.: 
White Mts., near McNarry P.O., 1-5 Aug. 1925, 1 d, 1 2, O.C. Pol- 
ing, ¢ genitalia made VI-8-35, A.G. Richards. Cochise Co.: Ash 
Canyon Rd., (1/2 mi W Hwy 92, Lot 4, 5100 ft. el.), 19 May 1981, 1 
2, June 1982, 2 2, 18 June 1980, 1 2, 13, 25 July 1980, 1d, 192, N. Mc- 
Farland; Cherry Canyon, near Cherry, 9 May 1959, R.F. Sternitzky, 
1 2; Chiracahua Mts., 3 d, 1 2, 26 June, H.G. Hubbard, 1 d, 2 2; 
Guadalupe Canyon, Peloncillo Mts., 4250, 19 June 1976, 5 d, 2 2, 
C.J.& S. Werner; Hereford, 2 d, 1 2°, C.R. Biederman; Huachuca 
Mts., 9d, 11 2, 6 genitalia made VI-5-35 A.G. Richards; Palmerlee, 
17 d, 24 2, 2 6 genitalia made VI-1-35, VI-5-35, A.G. Richards, 1-7 
July, 1 2, 8-14 July, 1 d, Aug. 1d, 1 2; Paradise, 15 3, 19 2, June 43 d, 
19 2, 3.6 genitalia made VI-5-35, 8-VI-1935 + H58 A.G. Richards, 
July 5 d, 5 2, Aug. 7 d, 9 2, Sep. 1 2; Southwest Research Station, 
Chiricahua Mts., 16 June 1963, 4 3, 1 2; SW Res. Sta., Chiricahua 
Mts., 5400’, 5-8 July 1964, D.R. Davis, 1 d, 1 2; SW Research Sta., 
Chiricahua Mts., 5 mi W Portal, 22 June 1956, O.L. Cartwright, 1 ¢, 
2 2; Wilgus, 3 d, 1 2; Ash Canyon, Huachuca Mts., 19 Oct. 1966, 1 d, 
22 Oct. 1966, 2 5, R.F. Stermitzky (CNC); Barfoot Park, Chiricahua 
Mts., 3 May 1934, G.H. & J.L. Sperry, 1 °? (AMNH); Carr Canyon, 
Huachuca Mts, 28 Sep. 1967, R.F. Sternitzky, 1 2 (CNC); Chiricahua 
Mts., Onion Saddle, 29 July 1996, Opler & Buckner, 1 2 (CSU); 
Cochise Stronghold, 3-5 Sep. 1965, J.T. Doyen, 1 d, 1 2 (UCB); 
Coronado Nat. For., Harshaw Cr., 7 mi SE Patagonia, 5 Aug. 1996, 
P. Opler, 1 ¢ (CSU); Coronado Nat. For., Upper Pinery Cyn., camp- 
ground, Chiricahua Mts, 30-31 July 1996, P. Opler & E. Buckner, 1 
é (CSU); Garden Canyon, Huachuca Mts., 26 July 1966, R.F. Stern- 
itzky, 1 6 (CNC); Guadalupe Cyn., 29 mi E Douglas, 15-16 Aug. 
1972, J.T. Doyen, 1 6 (UCB); Huachuca Mts., 30 May 1910, 1d, 1 
June 1910, 2 2, 5 June 1910, 1 3, L.R. Clemence (CNC); Huachuca 


146 


Mts., Ash Canyon Rd., 6 Aug. 1996, P.A. Opler, 1 ° (CSU); Miller 
Canyon, Huachuca Mts., 23 Aug 1966, R.F. Sternitzky, 1 6 (CNC); 
Parker Canyon, Huachuca Mts, 6 Sep. 1966, 1 d, 1 2, 16 Sep. 1966, 
1 6, R.F. Stemmitzky (CNC); Perilla Mts., 8 mi E Douglas, 29 Apr. 
1989, 1 d (UCB); Pinery Cyn., Chiracahua Mts., 6500’ , 6-7 Aug. 
1991, D. Rubinoff, YR. Hsu, J.A. Powell, 1 2 (UCB); Ramsey 
Canyon, 15 mi S Sierra Vista, Huachuca Mts., 6000’, 5 Aug. 1967, 1 
3,9 Aug. 1967, 1°, 15 Aug. 1967, 1d, 1°, Sternitzky (CNC); Ram- 
sey Canyon, Huachuca Mts., 11 May 1966 1 9, 7 June 1965, 1 9, 8 
Sep. 1967, 14, 1 2, R.F. Sternitzky (CNC); Rustler Park, Chiracahua 
Mts., 8500’, 3 July 1972, 1 9, 14 July 1972, 1 ¢, J. Powell, 2 Aug. 
1973, 1 9, J. Powell & S. Szerlip (UCB); Sierra Vista, 2 May 1966, 2 
2, 8 July 1967, 1 9, 23 July 1967, 1 2°, 17 Aug. 1967 1 9, 5 Sep. 1966, 
19, R.F. Sternitzky (CNC); Southwestem Research Station, Chiric- 
ahua Mts., 10 July 1962, C.W. Kirkwood, 1 d, 1 9 (CNC); Sunnyside, 
7 Sep. 1966, R.F. Sternitzky, 2 2 (CNC); SW Research Sta. of 
AMNH, Cave Creek Canyon, Chiracahua Mts., 5400 ft., 16 June 
1958, 1 °, J.M. & S.N. Burns, 28 June 1960, 2 d, 1 2, J.M. Linsley 
(UCB); SW Research Sta., 5 mi W Portal, 22 July 1959, 1 d, 4 Aug. 
1959, 1 ¢, E.G. Linsley, 24 July 1961, 1 9, J.M. Linsley (UCB); SW 
Research Sta., Chiracahua Mts., 9 Sep. 1958, E.G. Linsley, 2 6 
(UCB); Turkey Creek, Chiracahua Mts., 5600’, 1-2 Aug. 1986, J. 
Brown, 3 d, 1 2 (UCB): 4.5 mi SW Portal, 16-26 June 1981, 1 3, WS 
95-31 July, 1981, 1 °, 1-3 Sep. 1981, 1 5,192, K. & M. Cazier 
(SDNH); Chiricahua Mts., 3 July 1908,V.L. Clemence, 1 ¢ (SDNH); 
Chiricahua Mts., Portal, 17-19 Aug. 1979, D.K. Faulkner, 5 d, 8 2 
(SDNH); Douglas, Oct. 1961, 1 9 (SDNH); Dragoon Mts., Cochise 
Stronghold, 8 Aug. 1980, 1 ° (SDNH); Huachuca Mts., Ash Canyon 
Rd. (1/2 mi W Hwy 92, Lot 4, 1100 ft), 5 Apr. 1979, 1 °, 23, 27, 28 
May 1981, 2d, 4 2, 13 June 1979, 1 9, 25, June 1979, 1 d, 30 June 
1979, 1 2, 4, 8, 11, 14 July 1979, 42, 13 July 1980, 1°, 16 July 1981, 
1 4, 14,16, Aug. 1979, 3 3 25, 27, Aug. 1979, 2 3, 26 Aug. 1979, 1 9, 
27 Aug. 1981, 1 2, 1 Sep. 1981, 12, 10 Sep. 1980, 1 3, 6 Oct. 1980, 1 
3, 13 Oct. 1979, 1 3, 24 Oct. 1979 1 3, N. McFarland (SDNH); Rus- 
sler Park, Chiricahua Mts., 27, 29 June 1931, 2° (SDNH). Gila Co.: 
Christmas, 1 4, 1 2; 1902, O.C. Poling, 1 2; 15-30 June, O.C. Poling, 
19. Graham Co.: Safford, 8 Aug. 1964, J. Powell, 4° (UCB). Hemp- 
stead Co.: Hope, 22 July 1931, L. Knobel, 1 d, 3 genitalia slide I-5- 
32 A.G. Richards. La Paz Co.: Ehrenberg, 10 Aug. 1938, F.H. 
Parker, 1 °. Maricopa Co.: Phoenix, 2 9, Apr. 24-30, 1 2, 14 June, | 
2, J. Doll, 22 June, 1°, Collection Brklyn. Mus., 6, 12 Apr. 1978, 2 °, 
21, 23, 28, May 1978, 4°, 10 Aug. 1974, 1d, R. Wielgus; Rio Verde 
Mts., Phoenix, 9 1983, W. Barnes, 1 d, 2 2: Tempe, 5 Apr. 1920, E.V. 
Walter & H.L. Amold, 1 °; 24-30 July, 1 4; Scottsdale, Mt. Shadows, 
20-22 June 1978, M. Buegler, 1 6 (UCB). Mohave Co.: Hualapai 
Mts., 24-31 May, | 9; Pierce Ferry, 5-7 May 1967, D. Davis, 1 ¢. 
Pima Co.: Baboquivera Mts., Aug. 3 4, 1°; Baboquivera Mts., 5000 
ft., 1-15 June 1923, 9 d, 8 2, 1-15 June 1924, 21 d, 17 °,15-30 June 
1923, 110 6, 58 2, 8 d genitalia slide made VI-3-35 A.G. Richards, 
1-7 July 1923, 206, 16 9, 1-15 July 1923, 2 6,39, 1-15 July 1924, 15 
5,5 9, 15-20 July 1923, 1 2, 15-30 July 1923, 1 ¢, 15-30 July 1924, 
1d, 1-15 Aug. 1924, 2 3,12, 15-30 Aug. 1923, 1 3, 22, 15-30 Aug. 
1924, 1 2, 1-15 Sept. 1923, 3 9, 1-15 Sept. 1924, 1 3,42, 15-30 Oct. 
1923, 1 6, 1°, O.C. Poling; Redington, 16 4, 18 °, 2 d genitalia slides 
made VI-1-35 and VI-8-35 A.G. Richards; Santa Catalina Mts., 1 2, 
17 Aug., 3 °; Santa Rita Mts., ] Apr. 1940, 1 °, Aug. 1960, 1 d, 1 
Sep. 1960, 1 d, 2 2, K. Brown, 14, 15 June 1898, 1 6, 2 2, E.A. 
Schwarz, July 1 2, Coll. Brklyn. Mus., 26 July 1925, 1 d; Selis P.O., 
Indian Oasis, 1-10 May 1923, O.C. Poling, 1 d, 3 genitalia slide 
made VI-5-35 A.G. Richards; Tucson, May 22, 1 d, June 14,1 4,12, 
J. Doll, 1 2, B. Neumoegen; 3 mi N Ajo, 19 Feb. 1963, W.C. Cook, 1 
3 (CNC); Alamo Canyon, Ajo Mts., 22 Apr. 1947, G.H. & J.L. 
Sperry, 1 d (AMNH); Baboquivari Mts., 26 Apr. 1938, 28 Apr. 1938, 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


2°, G.H. & J.L. Sperry (AMNH); Baboquivari Mts., 5000 ft., 15-30 
June 1923, O.C. Poling, 1 6, 25 Apr. 1947, G.H. & J.L. Sperry, 1 ° 
(CNC); Blacksprings Campground, Madera Canyon, 10 July 1976, 
D. Whitman, 1 6 (UCB); Madera Canyon, 5-12 Sep. 1951, W. Ham- 
mer, 1 6 (CNC): Madera Canyon, Santa Rita Mts., 27 Aug. 1962, 1 
2, H.E. Milliron, 18 Sep. 1966, 1 9, 22 Sep. 1966, 1 3, C.W. Kirk- 
wood (CNC); Madera Canyon, Santa Rita Mts., 6 June 1968, 2d, 1 
2, P. Opler & J. Powell, 2 2, P. Opler, 3-5 Sep. 1969, 1 d, J. Powell 
(UCB); Molino Basin, Santa Catalina Mts., 4500 ft., 12 June 1968, 
J.M. & S.N. Bums, 1 6 (UCB); Organ Pipe Nat. Mon., 24 Mar. 1964, 
1 3, C.W. O'Brien, 1 3, C.N. Slobodchikoff (UCB); Tucson, 5 June 
1935, 1 2, 10 June 1935 1 2, Bryant, 30 June 1935, O. Watts, 1 d 
(CNC), 5 Mar. 1987, J. Reichel, 1 d (AMNH); Bates Well, 18 Nov. 
1939, 1 2 (SDNH); Madera Canyon, Santa Rita Mts., 4 Aug. 1980, 1 
2, 5 Aug. 1980, 1 9, C. Bruyea (SDNH); Santa Rita Mts., 14 Sep. 
1977, M. Plagens, 1 ° (SDNH). Pinal Co.: Superior, 5 June 1938, 1 
d, 6 genitalia USNM 46370, 15 June 1938, 1 °, S.E. Crumb; Oracle 
Junction, 3300, 9 May 1964, 1 2°, 12 May 1964, 1 2, W.C. Cook 
(CNC); Rancho Linda Vista, nr. Oracle, 4 May 1967, R.F. Stemitzky, 
1 6, 1 9 (CNC). Santa Cruz Co.: Madera Canyon, Santa Rita Mts., 
5100’, 10-26 July 1964, 2°, D.R. Davis, 5 Sep. 1956, 6 d, 3 9; 12 mi 
S Sonoita, Hidden Springs Canyon, 29 July 1966, R.F. Sternitzky, 1 
3, 1 2 (CNC); 5 mi SE Sonoita, 30 July 1966, R.F. Sternitzky, 1 d 
(CNC); Canelo, 22 Apr. 1966 1 °, 5 Sep. 1966, 1 d, 1 2, 12 Sep. 1966, 
2d, R.F. Sternitzky (CNC); Elgin, 14 Aug. 1966, 1 2, 4 Sep. 1966, 1 
2, R.F. Sternitzky (CNC); Madera Canyon, Rec. Area, Santa Rita, 30 
July 1991, P.A. Opler, 1 ¢ (CSU); Nogales, 12 July 1906, 1 d, 12 Aug. 
1906, 1 °, 14 Aug. 1906, 1 9°, 24 Aug. 1906, 1 oy, Dl Sep. 1906, 1 3,1 
2, A. Koebele (CNC); Patagonia, 30 Apr. 1910, 1 9, L.R. Clemence 
(CNC); Patagonia Mts., 7 mi SE Nogales, 1 Aug. 1991, P. Opler, Y.F. 
Hsu, D. Rubinoff, 1 ° (UCB); Sycamore Cyn., Hank Yank Springs, 
9 mi W Pena Blanca, 31 July 1991, P.A. Opler, 1 ° (CSU); Madera 
Canyon, 5 Aug. 1980, 1 d, 15 Aug. 1980, 1 °, 22 Aug, 1978, 3°, D.K. 
Faulkner (SDNH); Nogales, 9 Aug. 1977, D.K. Faulkner, 1 d, 1° 
(SDNH). Wasington Co.: 20 July 1966, R.L. Brown, 1 2 (CNC); 
Yavapai Co.: Congress Junction, 23 June 1938, 1 d, d genitalia 
USNM 46371, S.E. Crumb; Dewey, 16-23 June, 1 2; Douglas, 
16-23 May, 4 °, 8-15 June 3 2, 16-23 June 1 d, 1 °, 8-15 Aug., 1 9; 
Prescott, 16-23 June, 1 d, Bares Collection, July 14, 1 9, J. Doll; 4 
mi N Prescott, 22 May 1973, 1 6, 27 June 1973, 1 9, L.M. Martin 
(CNC); 5 mi N Prescott, 7 July 1973, L.M. Martin, 1 d (CNC); 
Granite Dells, 4 mi N Prescott, 8 Sep. 1971, L.M. Martin, 1 2 
(CNC); Mayer, 4800 ft., 26 Apr. 1961, R.F. Sternizky, 1 d (AMNH); 
Prescott, 29 May 1907, R.E. Kunze, 1 d (UCB), 2 Sep. 1907, 1 9, 8 
Sep. 1907, 1 °, 1909, 1 9, R.E. Kunze (CNC), 30 May 1907, R.E. 
Kunze, 1 6 (SDNH); July 1910, O. Bucholz, 1 2 (CNC); Camp 
Verde, 11 June 1941, 1 9 (SDNH). Yuma Co.: Planet Mine, Bill 
Williams R., 24-31 May, 1 5; Welton, 1-6 Mar. 1925, 1 d, 3 2, O.C. 
Poling, 13 June 1939, 1 2, L.L. Stitt; Yuma, 22 July 1925, 3 d, 6 2; 
Martinez Lake, 10 June 1961, C.A. Toschi, 1 6 (UCB), 1 Apr. 1910, 

°, 4 Apr. 1910, 1 d, L.R. Clemence (CNC), 9 Aug. 1941, W.P. 
Medlar, 1 ¢ (SDNH); Wellton, 14 Apr. 1938, F.H. Parker, 1 
(CNC). Bill Wms. Fork, July, F.H. Snow, 3 2 . No specific locality, 
1-7 May 7 d, 4 2. Southern, 5 d, 6 2. Poling, 1 6, 1 9, Oslar. Wash- 
ington Mts., B.P. Clark, 2d,3 2. CALIFORNIA: Imperial Co.: Dix- 
ieland, 1-15 Mar. 1922, 1 2, 15-30 Mar. 1922, 2 4,5 2, 15-30 Apr. 
1922, 1 d,1-15 May 1922, 2 6, O.C. Poling; 16 mi NW Westmore- 
land, 2 Nov. 1974, ]. Powell, 1 ¢ (UCB); Fish Creek Mts., 1 Apr. 
1955, D.F. Hardwick 1 d (CNC); Harpers Well, Mar. 1981, J. Smiley, 
1 2 (UCB); Carrizo, 10 Mar, 1928, 1 2, 10 June 1928, 192, C.C. Searl 
(SDNH); 5 Apr. 1941, W.P. Medlar, 1 2° (SDNH). Inyo Co.: 1-15 
Apr. 2d, 1 2;9 mi W Lone Pine, 19 July 1961, P.D. Hurd, J. Powell, 
1 5 (UCB); Furnace Creek, Death Valley, 12 Apr. 1957, R.L. 


VOLUME 56, NUMBER 3 


Langston, 1 6 (UCB); Shoshone, 9 Apr. 1962, R.W. Thorp, 1 2 
(UCB); Surprise Canyon, Panamint Mts., 24 Apr. 1957, J. Powell, 1 
2. (UCB); Tecopa, 7 June 1961, R.P. Allen, 1 3, 1 2 (UCB). Kem Co.: 
Taft, 28 Nov. 1942, 1 9 (CNC). Los Angeles Co.: Mt. Lowe, 8 July 
1924, H.G. Dyar, 1 2; Santa Catalina Island, 20 Oct. 1932, D.. Mead- 
ows, 1 d; 1 4, d genitalia made VI-2-35 A.G. Richards, May 1 °, Co- 
quillet; Pasadena, 16 June 1910, L.R. Clemence 1 6 (CNC); San 
Clemente Isl., Stone Biol. Sta., 4-6 Dec. 1981, J. Powell, 1 2 (UCB). 
Orange Co.: Warehouse, U.C. Irvine, 6 July 1969, 1 d (UCB). River- 
side Co.: Indio, 25 Feb. 1940, 3 d, 1 °, D. Meadows, 11, d genitalia 
slide made VI-2-35 A.G. Richards, 4 aes slide USNM 40524, 
12, 13, 14, May 1921, 8d, 3 9, » E. Piazza; 2 mi N. of Desert Center, 
31 Aug. 1946, C.I. Smith, 2 2 (UCB); 4 mi S Palm Desert, 2 July 
1963, R.L. Langston, 2 d, 2 ° ues), 5 mi NW Palm Spirngs, 20 
Sep. 1961, J.A. Chemsak, 1 2 (UCB); 6 mi N Desert Center, 31 Aug. 
1946, C.I. Smith, 2 ¢ (UCB); Chino Canyon, Palm Springs, 10 June 
1930, 1 2, 10 Apr. 1950, E.C. Johnston, 2 2 (CNC); Hopkins Well, 16 
Apr. 1958, J. Powell, 2 d, 3 9 (UCB); Indio, 24 Feb. 1955, 1 d, 20 
Apr. 1955, D.F. Harwick 1 d, 1 29 (CNC), 20 Apr. 1942, 1 9, 17, 20 
May 1942, 2°, 2 June 1942, 3 d, 19,5 June 1942, 1 2, 30 June 1949, 
1 2, W.P. Medlar (SDNH): painted Canyon, near Mecca, 27 Mar. 
1971, L. Orsak, 1 d (UCB); Palm Springs, 3 Apr. 1932, 1 d, 10 Apr. 
1934, 1 2 (CNC), 24 Jan. 1947, 1 2, 13 Oct. 1945, 1 2, C. Smith 
(UCB), 22 Mar. 1916, 1 2, 28 Mar. 1916, 1d CLL. Fox (SDNH); San 
Jacinto Mts., Pinyon Flat, 10 July 1967, J.W. Johnson, 2 ¢ (UCB); 
Thermal, 10 July 1956, M. Wassbauer, 1 2 (UCB); Thousand Palms, 
14 Feb. 1955, 1 9, 29 Apr. 1955 1 d, 1 9, J.E.H. Martin (CNC); San 
Bernardino Co.: Loma Linda, 12 May 1912, 1 d, d genitalia slide 
USNM 40523, G.R. Pilate, 8-15 Apr., 1 d, 1 9 - genitalia slide 
USNM 46322, 8-15 May, I 6, 16-23 io 90 oS 9 gentiealtea made VI- 
11-35 A.G. Richards, 8-15 June, 1 9; Coquillet; 10 mi NE Earp, 22 
Apr. 1960, J.R. Powers, 2 4, 1 2 (UCB); 29 Palms, 21 Apr. 1950, E.C. 
Johnston 1 2 (CNC); 6 air mi W Rice, dunes, 1 Apr. 1978, J. Powell, 
R. Dietz, 1 2 (UCB); 9 air mi S Baker, Zzyzx Springs, 20 Apr. 1977, 2 
d 4°, Chemsak & Powell, 21 Apr. 1977, 3 3, 2 2, Powell, 30 June 
1978, 1 2, Powell (UCB); Apple Valley, 19 May 1955, J.E.H. Martin 
1 2(CNC); Cronise Lake, 26 Apr. 1937, H.B. Leech 1 ° (CNC); Cro- 
nise Vy., 29 Apr. 1956, J. Powell, 1 ¢ (UCB); Joshua Tree Nat. Mon., 
15 mi S 29 Palms, 3600 ft., 19 Oct. 1966, D.F. Hardwick 4 4, 3 2 
(CNC); Wheaton Springs, Ivanpah Mts., 18 July 1947, C. Smith, 1 ° 
(UCB). San Diego Co.: Jacumba, ‘ 22 June 1924, H.G. Dyar, 1 6 
(USNM), 13 May 1978, 2 3, 26 Aug. 1979, 1 6 <} , J.W. Brown, 20 July 
1984, 3 2, 26 July 1980, 3 d, 9 Aug. 1980, 3 °, Faulkner & Brown, 4 
Aug. 1980, 1 6, 19(SDNED: Poon alley, 22 Sep. 1935, D. Mead- 
ows, 1 2; S. Felipe Val., 5 Sep. 1937, D. Meadows, 1 2; San Diego, 30 
Apr. 1924, H.G. Dyar, 1 6; 6 mi E Banner, 13 July 1963, J. Powell, 3 
3 (UCB); Borego, 24 Feb. 1940, 1 2 Woodcock, Mar. 1947, N. Crick- 
mer | 2 (CNC); Borego, 21 Apr. 1960, 1 2, J.F. Lawrence, 3 May 
1956, 1 d, J. Powell (UCB); Mt. Laguna, 4 mi NW, 5600’, 3 Nov. 
1966. D.F. Hardwick 1 3, 2 ° (CNC); Anza-Borrego Des. St. Pk., 
Grapevine Canyon, 25 Aug. 1986, Faulkner & Brown, | 2 (SDNH): 
Boulevard-Manzanita, 10 June 1979, 1 J, 3 June 1980, 1 2, R. Mess- 
ner (SDNH); NAS Miramar, Lep. Sur. Site 9, 11 July 1996, N. 
Broomfield, 1 d (SDNH); Ocenaside, 5 Aug. 1957, A.A. Lee, 1 6 
(SDNH); San Diego, 12 Oct. 1921, 1 2, June, 1 9, E. Piazza 
(SDNH); So. San Diego, 9 June 1978, 1 9 (SDNH). Stanislaus Co.: 
Del Rio, 26 Apr. 1922 1 2? (CNC). Werner Mts., 22 July 1922, 1 2, ex 
collection J.A. Comstock. La Puerta Valley, July 1911, G.H. Field, 1 
3 (UCB), July 1911, G.H. Field, 2 6, 8 9 (SDNH). Warners, Sep. 
1919, G.H. Field, 1 2 (SDNH). No specific locality, 1 °. COL- 
ORADO: Boulder Co.: Boulder, Silver Saddle Motel, 5500’. 8 June 
1961, M.R. McKay, 1 d (CNC). Fremont Co.: Canon City, 7 Aug. 
1973, G.F. Hevel, 1 3. Prowers Co.: Two Buttes Dam, 21 July 1990, 


147 


P.A. Opler, 1 d (CSU). KANSAS: Finney Co.: Garden City, F.B. Mil- 
liken, 1 °. Greenwood Co.: Eureka, 13 July 1954, E.L. Todd, 3 . La- 
bette Co.: Oswego, 23 May 1964, 1 d, 4 June 1965, 1 2, G.F. Hevel. 
MISSISSIPPI: Hinds Co.: Jackson, 19 June 1960, B. Mather, 1 ¢ 
(BM): Clinton, 23 Mar, 1975, B. Mather, 1 ¢ ' (BM). ). Jackson Co.: 
Ocean Springs, 4 Aug. 1921, R.P. Bamhart, 1 3. Rankin Co.: Pearl, 
14 July 1970, B. Mather, 1 ° (BM). MISSOURI: Benton Co.: War- 
saw,l1 May 1962, J.R. Heitzman, 1 °. NEBRASKA: Cherry Co.: 
Hackberry Lake, Valentine N.W.R., 28 June 1983, D.C. Ferguson, 1 
2. Platte Co.: Columbus, 24 June 1937, E.A. Froemel, 1 4. 
NEVADA: Clark Co.: Charlestown Mts., 1-15 July 1921, O.C. Pol- 
ing, 1 d; Las Vegas, 15 June 1905, 1 4, 2, 14, 20 June 1905, 3 2, T. 
Spalding; 16-23 Mar., 1 °, 16-23 May, 1 2°, 8-15 June, 3 d, 24-30 
June, 1 d, 24-30 July, 1 2. Nye Co.: Mercury, 4 Aug. 1965, 1 6. NEW 
MEXICO: Bernalillo Co.: 15 mi E Albuquerque, 3 Sep. 1975, La- 
fontaine & Bowen | 2 (CNC). Catron Co.: Gila Nat'l. Mon., 6000’, 4 
July 1964, D.R. Davis, 1 °. Chaves Co.: Roswell, Aug., Cockerell, 1 
2, 12 June, on grape, 1 ?, AG. Hammar. Colfax Co.: 2 mi E Ute 
Park, 15-18 July 1974, E.L. Todd, 1 2; Cimarron Canyon, Sangre de 
Cristo Mts., 7900 ft., 11 July. ee 1 d, E. & I. Munroe (CNC); Ra- 
ton, 6600 ft., 5 July 1962, 1 d ’. E.G. Munroe (CNC). Dona Ana 
Co.: Organ Mts., Finley an 5200 ft., 4 May 1979, R. Holland, 
1d, 1 2(AMNH). Eddy Co.: Carlsbad, 17 May 1950, E.C. Johnston 
1 2 (CNC); White City, 14 May 1950, 1 S, 15) May 1950, 1 °,H86 
E.C. Johnston (CNC). Gaudalupe Co.: Santa Rosa, 12 July 1951, 
A.K Wyatt, 1 2. Grant Co.: 2 mi SW Separ, 9 July 1964, J. Powell, 1 
° (UCB). Hidalgo Co.: Rodeo, 4 Sep. 1959, 1 6, D.D. Linsdale, 7 
Sep. 1959, 1 ¢, J.R. Powers (UCB). Lincoln Co.: Cedar Creek 
Campground, Ruidoso, 7000’, 28 July 1962, 1 4, 29 July 1962, 12, E 
& I. Munroe (CNC). Luna Co.: Deming, 8-15 July, 3 ¢, 16-23 July, 
7d, 1 9, 1 2. McKinley Co.: McGaffey, 23 July 1962, 1 5, E. & I. 
Munroe (CNC); McGaffey, Zuni Mts., 7500 ft., 22 rey 1962, 12, E 
& I. Munroe (CNC). Otero Co.: High Rolls, Sep. 1 6, 2 2, ¢ genitalia 
made VI-3-35 A.G. Tee 3$ 2; James Can., Sacramento Mts., | 
July 1940, D.G. Hall, 10 ¢, 9 2; Alamogordo, 10 May 1950, E.C. 
Johnston 12(CNC). poe) a sTucumeatl 25 June 1924, ].R. Hor- 
ton, 1 3. Rio Arriba Co.: 2 mi E Gobernador, 6700’, 20 Aug. 1971, 
D.F. Hardwick 1 9 (CNC). Sandoval Co.: Jemez Springs, 1 d; Frijoles 
Canyon, Bandelier Nat. Mon., 6050 ft, 18 July 1962, 1 4,12, E. & I. 
Munroe (CNC). Socorro Co.: Socorro, 27 July 1962, 1 °, E. Munroe 
(CNC); Gran Quivira Nat'l Mon., 6600’, 1-3 July 1964, 3 4,5 2, D.R. 
Davis. Wemple, 8 July 1939, 1 ¢, 11 July 1939, 3 2°, D. Meadows 
(USNM), 9 Sep. 1939, 1 6 (CNC). OKLAHOMA: Cleveland Co.: 
Norman, 18 May 1950, W.J. Reinthal 1 6 (CNC). Coe Co.: 
Witchata Nat. Forest, 11 June 1926, T.H. Hubbell 1 d, 2 9: Witchata 
Falls Nat. W. Ref., The Narrows, 10-18 July 1984, D. & M. Davis, 3 
2, Love & Carter Cos.: Lake Murray, 20 May 1950, W.J. Reinthal 1 
3, 22 (CNC). Murray Co.: Arbuckle Mts., 1 km W Turner Falls, 28 
[feme—2 July 1984, D. & M. Davis, 2 », Paine Co.: 1962, J.F. Reinert, 
1 3. TEXAS: Bexar Co.: San Antonio, 5 d, 3 °, 2 ¢ genitalia slides 
USNM 40525, 46326, d genitalia slide made XII-24-32 A.G. 
Richards, ¢ genitalia slide made VI-8-35 A.G. Richards, H. Schwarz, 
1 2, 1-7 Apr, 1 2, June 1899, 1 2, July 1899, 1 2 OC. Poling 
(USNM), 19 May 1939, Newman, | d , 1 2 (CNC). Blanco Co:: 
Blanco, 23 May 1963, A. & M.E. Blanchard, 1 2. Brazoria Co.: 
Churchill Bridge, 24 June 1962, A. & M.E. Blanchard, 1 ¢. Brewster 
Co.: Alpine, 2 ¢, 1928, 15-31 July 1926, 2 4, 2 d genitalia made VI- 
5-35 A.G. Richards, 8-14 Aug. 1926, 1 9, 2 genitalia made VI-12-35 
A.G. Richards, 1926, 2 d,1 2, 8-14 Mar. 1926, 1 2, 22-31 Mar. 1926, 
1 2, 1-7 Apr. 1926, 3 2, 8-14 Apr. 1926, 32 , 15-21 Apr. 1926, 4d, 4 
2, 22-30 Apr. 1926, 1 4 si, 7 May 1926, 2 12, 2 genitalia made VI- 
12-35 A.G. Richards, 8-14 May 1926, 5 152 May Wee, 12 7 
June 1926, 1 2, 6-14 July 1926, 1 d, 15-21 1 July 1926, 2d, 2 2, 22-31 


148 


July 1926, 2 d, O.C. Poling; Marathon, 7 June 1908, Mitchell & 
Cushman, 1 2 (USNM), 23 May 1950, E.C. Johnston, 1 ° (CNC); 
1-7 Aug., 2 3, 15-30 Aug., 5 3, 3 2, 15-30 Aug. 1926, 1 9, O.C. Pol- 
ing; Big Bend Nat. Park, 24 May 1950, E.C. Johnston 1 GS, Il @ 
(CNC); Hot Springs, Big Bend Nat. Park, 29 Apr. 1959, 1 ¢, M.R. 
MacKay, 6 May 1959, 1 °, L. Bottimer (CNC); Panther ise Big 
Bend Nat. Park, 19 May 1959, M.R. MacKay, 1 ° (CNC); The Basin, 
Big Bend Nat. Park, 4 May 1959, 1 4, 3°, 8 May 1959, 1 d, 2 9, 10 
May 1959, 2 2, 11 May 1959, 1 2°, 16 May 1959, 1 °, M.R. MacKay 
(CNC). Burnet Co.: San Antonio, Shovel Mt., June 1899, O.C. Pol- 
ing, 1 9, 2 genitalia slide USNM 46327; Shovel Mt., 2d, 2°, 8-15 
Apr. 16, 8-15 a: 2, 24-30 May, 1 2, 16-23 June 1 °. Cameron 
Co.: Brownsville, 2 3, Mar. 1911, 1d, May 1911, 2 2, June 1911, 3 2, 
G. Dorner, | Apr. 1999, ] 2, 25 Apr. 1928, 1 2, 31 May 1915, 2 8, 
M.M. High, 1 Apr. 1929, 1 9, 25 Apr. 1928, 2 9, 1, WB, DH, DB, Bel, ty, 
26, 27, 29 May 1928, 4 d, 19 2, 25 Sep. 1928, 1 °, FH. Benjamin, 
June 1911, 1d, 6 genitalia slide USNM 40521, G. Dorner, 12 Nov. 
1968, 1 d, A. & M.E. Blanchard, 1 July 1895, 1 °, Townsend, 1-7 
Aug. 1926, 1 2, O.C. Poling, 7 June 1904, 1 9, HLS. Barber; 
Browns ee Ranch, 1 2; San Benito, 7 2°, 1-7 Apr., 1d, 8-15 
May, 3, 19, 16-23 May, 58 d, 30 2, 2 d genitalia made VI-7- 35 A.G. 
Richards, d  ecoitelta made VI-9- 35 A.G. Richards, 2 2 genitalia 
made VI-11-35 A.G. Richards, 16-23 July 1 °, 8-15 Sept., 1 4, 1 9; 
Harlingen, 13 May 1958, J. Hunt, 2 ¢ (UCB). Chambers Co.: Black 
Jack Sprgs., 1 2. Cottle Co.: 8 mi NW Paducah, 1800, 23 Sep. 1968, 
2d, 24 Sep. 1968, 1 6, 1 °, D.F. Hardwick (CNC). Crane Co.: Crane, 
28 May 1950, E.C. Johnston 2 d, 3 2 (CNC). Culberson Co.: Sierra 
Diablo, 20 mi NNW Van Horn, 27, 29 May 1973, D.C. Ferguson, 5 
3, 4 2; Smith Cany., Guadalupe Mts., 5750’, 22 May 1973, D.C. Fer- 
guson, 1 2. Dallas Co.: Dallas, 22 May 1906, F.C. Pratt, 1 4, 
(USNM), 12 June 1942, CI. Smith, 1 9 (UCB): vic. Irving, 29 Apr. 
1984, 1 2, 6, 27 May 1984, 1d, 2 2, 28 May 1983, 1d, 12, 3, 23, 31 
May 1981, 3 d, 3 2, 1 June 1980, 12 d, 3 June 1981, 2 d, 10 June 
1983, 1 d, 10 June 1984, 1 d, 1 9, 22 June 1980, 1 9, R.A. Rahn; Irv- 
ing, 15 Mar. 1953, Woodcock, 1 6 (CNC). Duval Co.: San Diego, 20 
May 1895, bred from larva on Mesquite, E.A. Schwarz, 1 °. El Paso 
Co.: El Paso, 4000’, 19, 24 June, 2 ¢, 6 Aug., 1 2, J. Woodgate 
(USNM), 26 May 1964, J.E.H. Martin, 1 ° (CNC). Gonzales Co.: 
Ottine, Palmetto St. Pk., 12-13 Aug. 1963, P.J. Spangler, 1 °. Harris 
Co.: July, 1 °; Houston, Aug. 1916, 1 6. Hemphill Co.: cangie 
Wldlf. Mgt. Area Canadien, 11-12 July 1974, E.L. Todd, 1 4, 2 °. 
Hidalgo Co.: Donna, A.W. Nations, 1 °. Jeff Davis Co.: Davis Wee. 
Mount Locke, 6700’, 30 Aug. 1969, A. & M.E. Blanchard, 1 2; Davis 
Mtns. Resort, 5800’, 27 Apr. 1992, D.G. Marqua, 1 d (UCB); Ft 
Davis, 29 May 1959, 2 6, 30 May 1959, 3 2, 31 May 1959, 1 4, 5 @, 
M.R. MacKay (CNC). Kerr Co.: Kerrville, 3°, 11, 12 Apr. 1907, 1 
3,1 2, 30, 31 May 1906, 3 d, 6 2, d genitalia made VI-2-35 A.G. 
Richards, 1 June 1906, 1 d, 2 9, F.C. Pratt, May 1912, 1 d,1 2, H. 
Lacey, 16-23 Apr., 1 °, May 1903, 1 9, 17 May 1902, 12 (USNM), 5 
Apr. 1959, 1 2, 15 Apr. 1959, 1 d, R.H. Wigmore (CNC). Kimble Co.: 
Junction, 3 Apr. 1968, A. & M.E. Blanchard, 1 d, 14-17, June 1972, 
10 6, 9 2, D.C. Ferguson. La Salle Co.: Artesia Wells, 12, 13 June 
1972, D.C. Ferguson, 3 6; Cotulla, Crawford & Pratt, 2 3, 2 d geni- 
talia made VI-8-35 A.G. Richards. Menard Co.: Menard, 25 July 
1940, Hall, 2 3,1 2°. Nueces Co.:: ens Christi, May, 1 és Nueces 
River, 1 d, 26, 28, Apr. 1910, 2 d, F.C. Pratt, 22 June, 1 d, J. Doll. 
Pecos Co.: Ft. Stockton, 14 Ae. 1938, D. Meadows, 1 2. Presido 
Co.: Presidio, 20 July 1944, ex. Prosopis, 1 d, 3 °; Shafter, 4000’, 31 
May 1973, D.C. Ferguson, 1 d, 1 2, 16 Oct. 1973, 1 9, A. & M.E. 
Blanchard. Randall Co.: Palo Duro Canyon State Pk., 15 Apr. 1969, 
A. & M.E. Blanchard, 1 ¢. Reeves Co.: Pecos, 6 Sep. 1952, 2 °, 8, 9 
Sep. 1952, 2d, R. Leuschner (USNM), 18 May 1950, E.C. Johnston 
15,19 (CNC). Robertson Co.; Calvert, G.H. Harris, 1 2. San Patri- 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


cio Co.: Sinton Welder Wildlife Foundation (San Patricio), 24, 25 
May 1962, 14,19, 22 June 1962, 39, ? genitalia slide USNM 46376, 
22 Aug, 1962, 1d, 1 2, 13 Sep. 1963, 1d, 7 Oct. 1964, 2°, 2 genitalia 
slide USNM 46373, 13 Noy. 1963, 1 2, 2 genitalia slide USNM 
46374, A. &M.E. Blanchard. Terrell Co.: Sanderson, 27 Apr. 1959, 5 
3, 2°, M.R. MacKay, 28-29 Apr. 1959, 1 °, W.R.W. Mason (CNC). 
Travis Co.: Austin, 3 May 1897, H. Schwarz, 2 4, 6 genitalia made 
VI-4-35 A.G. Richards; Bee Cave, 4 Sep. 1962, 1 J, 1 9, A. & MLE. 
Blanchard, ¢ genitalia slide USNM 46372. Uvalde Co.: Sabinal, 1 
Apr. 1910.1 d, 1 2, 26 May 1910, 1 2, F.C. Pratt. Victoria Co.: Victo- 
ria, 22 May 1915, 1 d, 18 June 1911, 1 2, J.D. Mitchell. Walker Co.: 
Huntsville S’te. Park, 19 May 1963, 1 °, A. & M.E. Blanchard, 2 
genitalia slide USNM 46375. Webb Co.: Laredo, 1 Aug. 1938, D. 
Meadows, 1 ¢,1 °. Williamson Co.: Georgetown, 13 Apr. 1937, L.J. 
Milne, 1 ¢ (CNC). Belfrage, C.V. Riley, 1 2. 16 mi ESE Canyon, 
3000’, 26 Sep. 1968, D.F. Hardwick, 3 3, 1 2 (CNC). UTAH: Utah 
Co.: Vineland, 7 Aug. 1912, T. Spalding, 1 4. Washington Co.: St. 
George, 24-31 May, 1 9, 1-7 June, 1 6, 3 9, 8-15 June, 1d. 

Larval host. Prosopis sp. (Fabaceae). A female from the vicinity 
of Presidio, Texas in the USNM was reared. 

Distribution (Fig. 61). Southern Manitoba, south to Nebraska 
and east to Mississippi, west to California and south to central Mexico. 


Discussion. The type of B. deducta is the usual 
form the of male (Richards 1936a) (Fig. 25). The fe- 
males have three distinct phenotypes. One is the mac- 
ulate female that has all of the forewing pattern of the 
male, but is less distinct, and the median band and 
reniform spot are not as bright, but more infuscated 
with light brown (Figs. 26-27). The second is the im- 
maculate female that was described as B. pavitensis 
(Figs. 28-29). The third form,vulpina, is a female with 
the terminal half of the forewing with cinnamon 
ground color (referred to as “dull foxy red” by Ed- 
wards (1882)) (Fig. 30). Cirrhobolina incandescens 
Grote is a large female and the form albina is a light 
female (Richards 1936a). 

Collections indicate that B. deducta has a more mi- 
gratory pattern than other species of Bulia based on 
the specimen record from southern Manitoba, 
Canada. There is a definite southern limit to the distri- 
bution in Mexico, from a line extending from the vicin- 
ity of Monterrey and Linares west to El Salto and Du- 
rango. A single stray specimen from southeastern 
Chiapas, Mexico, is probably a migrant. 


ACKNOWLEDGEMENTS 


We express our gratitude to the 1999 Research Training Program 
at the National Museum of Natural History, Smithsonian Institution, 
Washington, D.C. Partial funding for this study was provided by the 
National Science Foundation Louis Stokes Alliances for Minority 
Participation (HRD9732790). We thank the following individuals for 
letting us borrow specimens under their care: J. Donald Lafontaine 
(CNC), Jerry A. Powell (UCB), David Faulkner (SDNH), Eric 
Quinter (AMNH), Manuel Balcézar Lara (UNAM), and Jorge F. 
Corrales (INBIO). We thank the following individuals who loaned 
specimens for this study: Daniel H. Janzen, “Unathy ersity of Pennsylva- 
nia, Philadelphia, Eric H. Metzler, Columbus, Ohio, and Bryant 
Mather, Clinton, Mississippi. For providing critical evaluation of the 
manuscript we thank David R. Smith and Stuart H. McKamey, Sys- 
tematic Entomology Laboratory, Washington, D.C. 


VOLUME 56, NUMBER 3 


LITERATURE CITED 


Barnes, W. & J. McDuNNouUGH. 1917. Check list of the Lepi- 
doptera of boreal America. Herald Press, Decatur, Illinois, ix + 
392 pp. 

Beur, H. 1870. Synopsis noctuidarum hucusque in California 
repertarum, Trans. Am. Ent. Soc. 3:23-28. 

Druce, H. 1889. Lepidoptera-Heterocera. In F. D. Godman and 
O. Salvin. 1881-1891. Biologia Centrali-Americana; or contri- 
butions to the knowledge of the fauna of Mexico and Central 
America. Zoology. Lepidoptera. Heterocera by H. Druce. Vol. 
1. London. Taylor and Francis. 490 pp., plates 1-64. 

Dyar, H. G. 1902 [1903]. A list of North American Lepidoptera 
and key to the literature of this Order of insects. Bull. USNM, 
no, 52, xix + 723 pp. 

Epwarbs, H. 1882. New species of Heterocera. Papilio 2:9-15. 

Farris, J.S. 1988. Hennig86 reference. Version 1.5. 

FRANCLEMONT, J. G. & E. L. Topp. 1983. Noctuidae. In R. W. 
Hodges, et al. (eds.), Check list of the Lepidoptera of America 
north of Mexico. E. W. Classey Limited and The Wedge Ento- 
mological Research Foundation. Cambridge, University Press. 
xxiv. 284 pp. 

GRIN DataBase. 2000. The Germplasm Resources Information 
Network (GRIN), GRIN Taxonomy. Agricultural Research 
Service, U.S. Department of Agriculture, Beltsville, Maryland. 
Website (http://vww.ars-grin.gov/npgs/tax/index.html). 

Grote, A. R. 1875. Descriptions of North American moths. Trans. 
Am. Ent. Soe. 5:113-118. 

HOLLAND, W. J. 1903. The moth book. A popular guide to the 
knowledge of the moths of North America. Doubleday, Page & 
Company, New York. 479 pp. 

JANZEN, D. H. 2000. Caterpillar rearing voucher databases for the 
Area de Conservacion (ACG) in northwestern Costa Rica. Web- 
site (http://janzen.sas.upenn.edu:591/database.htm). 

Kirsy, W. F. 1875. Lepidoptera. Zoological Record Insecta 
12:403—-467. 

Lipscoms, D. 1994. Cladistic analysis using Hennig§86, version 1.5. 
George Washington University, 122 pp. 

McDunnoucu, J. 1938. Checklist of the Lepidoptera of Canada 
and the United States of America. Part 1. Macrolepidoptera. 
Mem. South. Calif.Acad. Sci . 275 pp. 

Morrison, H. K. 1875. List of a collection of Texas Noctuidae, 
with descriptions of the new species. Proc. Bos. Soc. Nat. Hist. 
17:209-221. 

Nye, I. W. B. 1975. The generic names of moths of the World. Vol. 
1. Noctuoidea (part): Noctuidae, Agaristidae, and Nolidae. Lon- 


149 


don. Trustees of the British Museum (Natural History), Publi- 
cation Number 770, 568 pp. 

POOLE, R. W. 1989. Lepidopterorum Catalogus (New Series), Fas- 
cicle 118, Noctuidae. E. J. Brill and Flora and Fauna Publica- 
tions, Leiden. xii + 1313 pp. 

POOLE, R. W. & P. GENTILI (EDS.). 1996. Nomina Insecta Nearc- 
tica. A check list of the insects of North America, Vol. 3: 
Diptera, Lepidoptera, Siphonaptera. Entomological Informa- 
tion Services, Rockville, Maryland. 1143 pp. 

Ricuarps, A. G. 1933. Comparative skeletal morphology of the 
noctuid tympanum. Entomol. Am. 13:1-84. 

. 1936a. A revision of the noctuid genus Bulia Walker 

(=Cirrhobolina Grote) (Lipidoptera) [sic]. Ann. Ent. Soc. Am. 

29:431-437, 

. 1936b. A generic synopsis of the Phoberia-Melipotis Dras- 

teria-Boryza series of Erebinae (Lepidoptera, Noctuidae). Rev. 

Entomol. 6:338—374. 

. 1939. A revision of the North American species of the 

Phoberia-Melipotis-Drasteria group of moths (Lepidoptera, 

Phalaenidae). Entomol. Am. 19:1—99. 

. 1941. The genus Bulia Walker in Mexico and Central 
America (Lepidoptera, Phalaenidae). Allan Hancock Pacific Ex- 
peditions 5:255-267. 

SIMPSON, B. J. 1988. A field guide to the Texas trees. Texas Monthly 
Press, Austin, Texas. 372 pp. 

SMITH, J. B. 1891. List of Lepidoptera of boreal America. Philadel- 
phia. P. C. Stockhausen. v + 124 pp. 

. 1893. A catalogue, bibliographical and synonymical, of the 
species of moths of the Lepidopterous superfamily Noctuidae, 
found in boreal America. Bull. USNM, no. 44, 424 pp. 

STRECKER, F. H. H. 1900. Lepidoptera, Rhopaloceres and Hetero- 
ceres, Indigenous and Exotic. Supplement 3. [No publisher 
given]. 37 pp. 

WALKER, F. [1858] 1857. List of Specimens of Lepidopterous In- 
sects in the collection of the British Museum. Vol. 13. Edward 
Newman, London. pp. 983-1236. 

. 1858. List of the specimens of lepidopterous insects in the 

collection of the British Museum. Vol. 15. Edward Newman, 

London. pp. 1521-1888. 

. [1866] 1865. List of the specimens of lepidopterous in- 

sects in the collection of the British Museum. Vol. 34. Edward 

Newman, London. pp. 1121-1533. 


Received for publication 8 January 2001; revised and accepted 12 
January 2002. 


150 


bo 


10. 


11. 


13}, 


14. 


15. 


16. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


APPENDIX |. Characters and character states used for phylogenetic analysis of Bulia; 0 = plesiomorphic state, 1-3 = apomorphic states. All 
characters were treated as ordered, with Figure numbers illustrating those character states. 


Head 

Head 

Hindwing 
Eighth segment 
Eighth segment 
Uncus 


Valva 


. Valva 


Sacculus 


Sacculus 


Juxta 


Aedoeagus 


Vesica 


Coremata 
Genital plate 


Median prong 


0 


2 


9 


74 


0—vertex of male without an elongate median projection 


1—-vertex of male with an elongate median projection (Fig. 3) 


0—anal lunule white 
1—anal lunule yellow 
0—tergum not reduced, mostly sclerotized 


0—labial palpus third segment not extending above vertex 
1—labial palpus third segment extending above vertex 


1—tergum reduced to a narrow X-shaped tergite, mostly membranous (Fig. 31) 


0—sternum not reduced, mostly sclerotized 


1—sternum reduced to a wine glass shaped sternite, remainder membranous (Fig. 32) 


0—bearing long bifurcate setae 


1—long bifurcate setae absent 


0—clasper present 


1—clasper absent 


0—clavus present 


1—clavus absent 


lacking large spines 
1—with 2 large spines (Figs. 44-45, 47-48) 
2—with 1 large spine (Fig. 46) 


1—fused to the 7th sternum 


0—absent 
1—short, width greater than height (Figs. 49-51, 54-56) 
—long, width less than height (Figs. 52-53, 57-58) 


APPENDIX 2. Data matrix for 2 outgroups, Melipotis (designated 
outgroup) and Drasteria, and species of Bulia. Character numbers 
refer to those in Appendix 1. 


0—lacking an expanded costal lobe 

1—costal lobe small, triangular (Figs. 34-35) 

2—costal lobe large, apex slightly bifurcate (Fig. 36) 
3—costal lobe large, apex truncate to produced (Figs. 37-38) 
apex not differentiated 
1—apex not extending beyond cucullus 

apex extending beyond cucullus, short (Figs. 34-35) 
3—apex extending beyond cucullus, elongate (Figs. 36-38) 
0—dorsal margin sightly concave 
1—dorsal margin U-shaped (Figs. 34-37 and 39-42) 
2—dorsal margin V-shaped (Fig. 38 and 43) 
0—external spiculations absent 

1—external spiculations present (Figs. 47-48) 

0. 


0—if present, arising from base of valva 
1—arising from base of Sth tergum (Fig. 33) 
0—not fused to the 7th sternum 


Melipotis 


Drasteria 
confirmans 
mexicana 


schausi 


similaris 


deducta 


12345 67890 


00000 00000 
01000 10001 
aLaL@abal  dhababatA) 
ALababalal ababalalZ) 
abababal ab al alas) 
aaLILakal aaa) 
aatast ILLS 3} 


ababacalat 
12345 


00000 
00000 
10111 
10111 
AL@)AabaL 
aL aL al aL aL 
Balabalal 


I Ww Pep So oS 


Journal of the Lepidopterists’ Society 
56(3), 2002, 151-161 


FRANCIS H. SNOW’S NEW MEXICO BUTTERFLY COLLECTIONS 


STEVEN J. CARY 
202 Solana Drive, Santa Fe, New Mexico 87501, USA 


RICHARD HOLLAND 
1625 Roma NE, Albuquerque, New Mexico 87106, USA 


ABSTRACT. Francis H. Snow laid the foundation for scientific butterfly study in New Mexico. From his faculty post at the University of Kansas 
he led six natural history collecting expeditions into New Mexico between 1880 and 1894. These campaigns produced one butterfly species new to 
science, more than 90 state records, useful summary reports and labeled reference specimens. Butterflies collected on his first and last New Mex- 
ico expeditions are published here for the first time. Eighty labeled specimens from these six expeditions remain in the collections at the Snow En- 


tomological Museum at the University of Kansas. 


Additional key words: naturalist, natural history, collecting expedition, entomologist. 


There was little systematic Lepidoptera study in 
New Mexico before the expeditions of entomologist 
Francis Huntington Snow. With so little known about 
its insect fauna, New Mexico must have seemed to 
Snow like an ideal destination. He was the first to con- 
duct extensive butterfly collecting in New Mexico, the 
first to document detailed results in published reports, 
and the first to adequately label New Mexico speci- 
mens for future use. Snow's New Mexico contributions 
were first documented in published reports listing but- 
terflies collected on expeditions in 1881, 1882 (Snow 
1883), 1883 and 1884 (Snow 1885), which stood un- 
questioned for nearly 100 years. 

It was the late 20th century before knowledge of the 
New Mexico butterfly fauna advanced to the point 
where some of Snow’s published reports could be criti- 
cally examined. In 1978, M. Toliver sought to confirm 
or refute certain inconsistencies by examining speci- 
mens in the collections at the Snow Entomological 
Museum (SEM) at the University of Kansas (UK). To- 
liver’s determinations of dubious Snow reports helped 
consolidate historic knowledge of New Mexico and 
Rocky Mountain butterflies (Ferris & Brown 1980, 
Cary & Holland 1994, Toliver et al. 1994). 

Snow published collection reports for New Mexico 
expeditions from 1881 through 1884 (Snow 1883, 
1885). Snow (1883) briefly mentioned an 1880 New 
Mexico expedition, but implied that it produced no 
butterflies. He also mounted an expedition to New 
Mexico in 1894, for which he published Coleoptera 
collections (Snow 1907) but not Lepidoptera. Review 
of biographical material and re-examination of SEM 
collections in 1998, 2000 and 2002 confirmed that 
Snow collected butterflies in New Mexico in 1880 and 
1894 and revealed details about those expeditions. Re- 
search also showed that Snow's expeditions of 1881 
through 1884 produced more species than indicated in 


his reports. Previously unreported collections are de- 
scribed below. 

In this project, the authors profited from several 
sources of biographical information. Snow’s biogra- 
pher (Hyder 1953) listed all of Snow’s scientific expe- 
ditions including his 1880 trip to Santa Fe Canyon and 
the 1894 trip to the Magdalena Mountains. Following 
Snow's death in 1908, Snow’s colleagues eulogized him 
as a teacher, scientist and collector. Miller (1909), 
Marvin (1909), Stevens (1909) and Dyche (1909) pro- 
vided valuable information and insights into Snow's 
character and methods. Snow belatedly published 
combined Coleoptera results from his six New Mexico 
collecting expeditions (Snow 1907). It is worthwhile 
now to do the same for his butterflies. 


FRANCIS HUNTINGTON SNOW 


Born in 1840 in Fitchburg, Massachusetts, Francis 
H. Snow was raised with a strong work ethic and reli- 
gious faith. His father was a staunch abolitionist who 
gave haven to escaped slaves on the Underground 
Railroad. Snow matured as the nation considered 
statehood for Missouri and Kansas; the Snows worked 
to keep Kansas slave-free. Snow was valedictorian of 
the Williams College class of 1862. Toward the end of 
the Civil War, he provided spiritual and hospital sery- 
ices for sick, wounded and dying soldiers. 

Snow was classically trained and socially motivated, 
a practiced preacher and gifted educator. In 1866, five 
years after Kansas was admitted to the Union as a free 
state, Snow accepted a position with the newly 
founded University of Kansas in Lawrence. As one of 
the first three faculty members, he helped recruit stu- 
dents and assemble buildings, faculty and curricula. In 
1868 Snow returned to Massachusetts, married Jane 
Aiken and brought her to Lawrence. He was devoted 
to his family, which eventually included six children. 


Santa Fe 
CA 
Las Vegas 


4 
Albuquerque 
| Abuaverave # 


Magdalena@ e 
° 
*, Socorro 


Silver City® 


Fic. 1. Snow’s New Mexico expedition locations and railroad 
lines ca. 1890. 


Most of Snow’s collecting excursions were family af- 
fairs and he corresponded with his wife when circum- 
stances prevented her participation. 

Snow taught every branch of natural science at UK. 
He had to teach himself first because his classical edu- 
cation did not include the many new scientific sub- 
jects. During his career Snow published more than 
150 papers and reports. In 1890 Snow was named 
Chancellor of the UK. He served until 1901 when the 
strain of work and the untimely death of his first son 
forced him to cut back. Snow found solace and 
strength in the outdoors and, until his death in 1908, 
Snow did what he loved best—collecting, working 
with specimens, and teaching students about nature. 


Collecting Expeditions 


During his 42 years at UK, Snow lead 26 collecting 
expeditions to Kansas, Colorado, Texas, New Mexico 
and Arizona (Hyder 1953:279-280). For these expedi- 
tions, Snow sought out regions that offered opportuni- 
ties for naturalists to discover and study new flora and 
fauna. Dyche (1909:40) explained that Snow’s primary 
objective during these expeditions was “to make col- 
lections of insects, although he was interested in nearly 
every branch of natural history, and encouraged his as- 
sistants and students to collect birds, mammals, rep- 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


tiles, fishes and plants, as well as specimens of rocks, 
minerals and fossils.” 

Each expedition produced thousands of specimens 
(Miller 1909:24), which Snow examined upon his re- 
turn. Varieties not represented in the UK collection 
were sent to specialists for determination. Specimens 
that enhanced the breadth of the UK collection ulti- 
mately were retained, but thousands of others ended 
up in the duplicate collection. Snow treated duplicates 
as assets to be sold or traded with collectors in the U.S. 
and Europe in return for other desirable specimens 
(Dyche 1909:44). Although his expeditions produced 
hundreds of insect species that were new to science, it 
appears that Snow himself described none (Dyche 
1909:44). 


1880: Santa Fe Canyon, Sangre de Cristo Mountains 


The 1880 UK expedition came by railroad (Fig. 1) 
to Santa Fe, the capital of New Mexico Territory 
(Snow 1881b). Snow brought his family (Jane, 11-year 
old son Willie and 10-year old daughter Mattie) and 
two student assistants, Louis L. Dyche and Annie E. 
Mozley. The group was in the field from August 6 to 
September 3. From their camp in the canyon of the 
Santa Fe River, they collected in the southwestern part 
of the Sangre de Cristo Mountains in Santa Fe County. 

Confusion clouded the correct location of Snow’s 
1880 base camp. Snow first described it as “in Santa Fe 
canon” “eight miles from the city of Santa Fe, at an el- 
evation of 7,000 feet” (Snow 1881b:67), then later as 
“eight miles north of Santa Fe” (1907:165). Dyche 
(1909:42) stated it most clearly: “Camp was located 
eight miles up the cafion from Santa Fé.” Starting 
from the City at 7000 feet elevation on the Santa Fe 
River, eight miles up-canyon took Snow eastward to 
about 8000 feet elevation. 

Arriving in Santa Fe in 1880, Snow found a land- 
scape occupied by Spanish colonists almost continu- 
ously since ca. 1700. He traveled up the Santa Fe 
River along a well-worn trail first used by Indians, then 
by European settlers driving livestock to summer pas- 
ture (Fauntleroy 1999). Arrival of the transcontinental 
railroad in 1880 opened Santa Fe to nationwide eco- 
nomic markets, prompting local farmers to expand 
their herds. One year later near Snow's camp, the City 
constructed McClure Reservoir, which still remains 
part of the City’s public water supply system (Faunt- 
leroy 1999). 

The published report for the 1880 expedition listed 
273 species of Coleoptera, but no Lepidoptera (Snow 
1881b). Instead, Snow remarked on the “conspicuous 
scarcity in Santa Fe canon . . . of the Lepidoptera, both 
diurnal and nocturnal,” which he attributed to “nu- 


VOLUME 56, NUMBER 3 


TABLE 1. Butterflies collected during Snow’s 1880 expedition to 
Santa Fe Canyon, Santa Fe County, New Mexico. SEM = Snow En- 
tomological Museum at the University of Kansas. 


Number of SEM 


Taxon specimens Importance 
Erynnis pacuvius 1 
(Lintner, 1878) 
Apodemia nais 1 Ist NM specimen 
(W. H. Edwards, 1876) 
Speyeria hesperis i Ist NM specimens 


(W. H. Edwards, 1864) 


merous flocks of sheep and goats, which destroy the 
food plants of these insects” (Snow 1881b:68). 

Toliver’s 1978 review of SEM specimens revealed 
the first butterflies from this expedition (Table 1). He 
found two specimens of Speyeria hesperis (W. H. Ed- 
wards) and one of Erynnis pacuvius (Lintner) labeled 
from Santa Fe Canyon in August 1880. The senior au- 
thor discovered a similar specimen of Apodemia nais 
(W. H. Edwards) while researching collections of the 
National Museum of Natural History (NMNH) in the 
1980s. Recent review of the SEM collection revealed a 
second A. nais specimen with the 1880 label. Each 
specimen from this expedition is recognized by a type- 
set label reading “S. Fe Canon, N. M., 7000 ft., Aug., 
1880 (Snow).” 


1881: Water Canyon, Magdalena Mountains 


Snow returned to New Mexico in August 1881. 
Prompted, perhaps, by the previous year’s disappoint- 
ment, he ventured farther from civilization into wilder 
country, to Water Canyon in the northeast part of the 
Magdalena Mountains, Socorro County (Fig. 1). 
Snow's 1881 crew included son Willie, Mr. Dyche and 
physics professor H. S. S. Smith. 

As with much of south-central New Mexico at that 
time, the original Apache inhabitants still clung tena- 
ciously to their lands, which included the Magdalena 
Mountains. The prospect of travel there may have 
given the Snow household reason for pause. This ex- 
pedition began routinely, as Snow wrote to Jane on 
August 11: “... I was never before so far away from 
the railroad in camping out. We are 25 miles west of 
Socorro, New Mexico, so that it is entirely out of the 
question for any of us to go to town as we did at Santa 
Fe... Our camp is about 2 miles from the entrance to 
the cafion, near the log house of one John Smith who 
has lived here 8 years, during which time not a single 
Indian has entered the cafion . . . Willie is having a fine 
time and says he is glad he did not stay East. We both, 
however, miss you and the little girls sorely. . . Willie 
and I are getting lots of fine butterflies & other insects 


153 


& Dyche is putting up plants of which there are many 
new and beautiful species.“ 

Two days later, Snow's idyll was shattered when a 
messenger, “who showed a bullet hole in his hat,” in- 
formed the group that the local Apaches had become 
hostile. Snow’s group assembled at a cabin with some 
miners and kept watch for three days (Hyder 
1953:158-159). The group then headed for the safety 
of Socorro on a journey described by Snow: “We were 
asked to leave all our belongings in order to go down 
the cafion double-quick, but I could not see how a 
dozen or so of light cigar boxes filled with my bugs 
could endanger our march, so despite the wishes of 
the others I carried with me thirteen boxes of my 
bravest [finest] insects. I tied them up so I could get 
my arms around them and, putting my Boston Win- 
chester over my shoulder, and with little Will, my Le 
clinging to my sleeve, we started for Socorro... On 
the way down we found pools of blood where men had 
been murdered, and their wagons looted. Our team- 
ster was one of the victims. A posse of 100 men com- 
ing out from Socorro met us and with them we contin- 
ued our march down the cafion.” 

The 1881 expedition retreated to areas that were 
safer, but less desirable for collecting. They spent 
three weeks near Socorro, one day at Deming and a 
week near Pecos (Dyche 1909:42). Snow (1883:36) re- 
ferred his specimens to W. H. Edwards for determina- 
tion. This troubled expedition produced more than 
two dozen species of butterflies from Water Canyon 
(Table 2) and one from Deming, but none from So- 
corro or Pecos (Snow 1883). Each Water Canyon spec- 
imen in the SEM was identified based on a single 
typeset label stating: “Water Canon, N. M., 5000 ft., 
Aug. ‘81, F. H. Snow.” 

1881 is the first year for which Snow’s reports can be 
compared against extant specimens. Allowing for 
nomenclature changes, they match pretty well. The 
few discrepancies are understandable after more than 
a century of taxonomic progress. Snow's report of E. 
funeralis (Scudder & Burgess) probably was the simi- 
lar Erynnis tristis tatius (W. H. Edwards), which was 
not described until 1882 (Ferris & Miller 1980:21) but 
for which there is an 1881 SEM specimen. He also re- 
ported Erynnis martialis (Scudder), which is impossi- 
ble (see Cary & Holland 1992[1994]). He probably 
had E. afranius (Lintner) based on size, markings, 
habitat and voltinism, but this remains speculation un- 
til a specimen is located. The final 1881 mystery con- 
cerns Brephidium exile (Boisduval). Snow (1883) re- 
ported it only from Deming, but the 1881 SEM 
specimen has a Water Canyon label. 

Snow’s labeled elevation represented the mouth of 


154 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


TABLE 2. Butterflies collected during Snow's 1881 expedition to Water Canyon, Magdalena Mountains, Socorro County, New Mexico. 


Taxon 


Erynnis pacuvius (Lintner, 1878) 
Erynnis tristis (Boisduval, 1852) 
ssp. tatius (W. H. Edwards, 1882) 
?Erynnis afranius (Lintner, 1878) 
Pholisora catullus (Fabricius, 1793) 
Piruna pirus (W. H. Edwards, 1878) 
Pterourus multicaudatus (W.F. Kirby, 1884) 
Pterourus rutulus (Lucas, 1852) 
Zerene cesonia (Stoll, 1790) 
Phoebis sennae (Linnaeus, 1758) 
ssp. eubule (Linnaeus, 1767) 
Eurema mexicana (Boisduval, 1836) 
Eurema nicippe (Cramer, 1779) 
Nathalis iole Boisduval, 1836 
Hypaurotis crysalus (W. H. Edwards) 
Brephidium exile (Boisduval, 1852) 
Everes amyntula (Boisduval, 1852) 
Plebejus lupini (Boisduval, 1869) 
Euptoieta claudia (Cramer, 1775) 
Chlosyne lacinia (Geyer, 1837) 
ssp. crocale (W. H. Edwards, 1874) 
Phyciodes pictus (W. H. Edwards, 1865) 
Polygonia gracilis Grote and Robinson, 1867 
ssp. zephyrus (W. H. Edwards, 1870) 
Nymphalis milberti (Godart, 1819) 
Vanessa virginiensis (Drury, 1773) 
Vanessa cardui (Linnaeus, 1758) 
Limenitis weidemeyerii (W.H. Edwards, 1861) 
Adelpha bredowti Geyer, 1837 


Reported by Number of 
SEM! specimens 


Snow (1883) as 


Thanaos Pacuvius Lintn. 


Thanaos funeralis Scud.-B. ] 
Thanaos Martialis Scud. 

Pholisora catullus Cram. 

Pholisora Pirus Edw. 1 
Papilio Daunus Edw. 

Papilio Rutulus Bd. 

Colias Caesonia Stoll 


Callidryas Eubule L. 

Terias Mexicana Bd. 

Terias Nicippe Cram. 

Nathalis Iole Bd. 

Thecla Crysalus Edw. 

not reported 1 
Lycaena Amyntula Bd. 

Lycaena Acmon West-Hew. 

Euptoieta Claudia Cram. 


Synchloe Crocale Edw. 
Phyciodes picta Edw. 


Grapta Zephyrus Edw. 1 
Vanessa Milberti Godt. 

Pyrameis Huntera Drury 

Pyrameis cardui L. 

Limenitis Weidemeyerii Edw. 

Heterochroa Californica But. 1 


Importance 


Ist NM specimen 
Ist NM report 
Ist NM report 
lst NM specimen 
Ist NM report 
Ist NM report 
Ist NM report 


Ist NM report 
Ist NM report 
1st NM report 
Ist NM report 
Ist NM report 
Ist NM specimen 
Ist NM report 
Ist NM report 
_Ist NM report 


Ist NM report 
Ist NM report 


Ist NM specimen 
Ist NM report 
Ist NM report 
Ist NM report 
Ist NM report 
Ist NM specimen 


Cyllopsis pertepida (Dyar, 1912) Neonympha Henshawii Edw. 2 1st NM specimens 
Cercyonis meadii (W. H. Edwards, 1872) Satyrus Meadii Edw. 1st NM report 
Danaus plexippus (Linnaeus, 1758) Danais Archippus Fab. 1st NM report 
Danaus gilippus (Cramer, 1775) Danais Berenice Cram. Ist NM report 


'SEM = Snow Entomological Museum at the University of Kansas. 


the canyon rather than the whole area sampled, as it 
did for Santa Fe Canyon in 1880. Water Canyon 
crosses the 5000-foot contour in open desert grass- 
land, but Snow camped two miles above the canyon 
mouth at about 8000 feet elevation. In addition to 
SEM specimens from this expedition, the American 
Museum of Natural History (AMNH) appears to have 
three specimens of Cyllopsis pertepida (Dyar), cited 
by Miller (1974:72). 


1882: Gallinas Canyon, Sangre de Cristo Mountains 


In 1882 Snow traveled to Gallinas Canyon, which 
begins in the southern portion of the Sangre de Cristo 
Mountains and descends southeastward through Las 
Vegas, in San Miguel County (Fig. 1). Arrival of the 
railroad in 1879 changed Las V egas from a quiet farm- 
ing village to a busy railroad town. Construction of a 
resort hotel near the hot springs in 1880 quickly made 


Las Vegas an international destination for consump- 
tives and other health seekers. 

The 1882 group, consisting of Snow, his family and 
three university students: Mr. Dyche, Mary Dyche, 
and Mr. W. W. Russ, produced more butterfly records 
than Snow's other New Mexico adventures. In Gallinas 
Canyon, Snow found a “naturalist’s paradise” com- 
pared to the 1880 expedition and a safe haven com- 
pared to the expedition of 1881. “The occupancy of 
the mouth of the cafion by hotels and bathing-houses 
of that famous sanitarium, has proved an effectual bar- 
rier against the entrance of troublesome ruminants, 
and the entomologist and botanist is able to obtain the 
choicest of his favorite objects of study in delightful va- 
riety and perfection” (Snow 1883:35). 

After nine weeks of collecting, Snow (1883) re- 
ported about 50 butterfly taxa (Table 3). From this ex- 
pedition, the SEM retains 30 specimens representing 


VOLUME 56, NUMBER 3 


27 species, including three not given in Snow’ report. 
Most specimens bear typeset labels stating: “Near Hot 
Springs, Las Vegas, N.M., 7000 ft., July, 82, F. H. 
Snow.” Others give the date of collection as “Aug. 82.” 
Two reported species, Pieris sisymbrii (Boisduval) and 
Anthocharis sara Lucas, fly only in spring; an SEM 
specimen of the former has “April” inked in longhand 
over the typeset “July.” Archival materials suggest that 
Snow had an associate in Las Vegas who could have 
obtained these specimens in the proper season. 

Once again, Snow’s party explored a range of eleva- 
tions not reflected on the specimen labels. Collecting 
at about 6800 feet elevation in foothill canyons near 
the Hot Springs probably produced Upper Sonoran 
Zone species like Amblyscirtes aenus W. H. Edwards. 
Gallinas Creek and its tributaries continue upstream 
from there, climbing gradually for more than 20 miles 
through Transition Zone into Canadian Zone habitats. 
Snow's group likely traveled this route, visiting eleva- 
tions near 10,000 feet elevation to collect Speyeria 
mormonia Boisduval, Oeneis chryxus (Doubleday & 
Hewitson) and Erebia epipsodea Butler. 

There are a few mismatches between SEM speci- 
mens and Snow (1883) reports (Table 3), primarily 
among the Hesperiidae. Snow reported Thorybes ba- 
thyllus (J. E. Smith), but the SEM specimen so labeled 
is actually T: pylades (Scudder). There also is confusion 
among the Erynnis spp., which were not well known at 
the time. An SEM specimen of E. horatius (Scudder & 
Burgess) is mislabeled as E. martialis (Scudder), yet 
neither can be linked conclusively to the reported E. 
juvenalis (Fabricius), which is erroneous. Among the 
SEM specimens labeled to the 1882 expedition are 
three that Snow (1883) did not report (Table 3). 

-Snow collected the type series of Hesperia viridis 
(W. H. Edwards) in 1882 during a period when skip- 
pers once lumped with Old World Hesperia comma 
(Linnaeus) were being differentiated and described. 
Amid this taxonomic instability, Snow (1883) reported 
collecting “Pamphila Comma n. var.” Edwards (1883) 
described the male “sent me by Prof. Snow and taken 
by him at Los Vegas [sic], N. M., in 1882” and called it 
“var. Viridis.” This was the only new taxon described 
using butterflies collected in New Mexico by Snow. 

Two specimens found by F. M. Brown in the 
AMNH in the 1970s may be traceable to the 1882 ex- 
pedition. A single specimen of Euphyes vestris (Bois- 
duval) appears to have the distinctive Snow label for 
that expedition (Toliver et al. 1994:112). A specimen of 
Hemiargus ceraunus (Fabricius) is similarly labeled, 
but is dated “Sept.” (Toliver et al. 1994:258). Each has 
a label indicating that it came to the AMNH via the 
Hulst collection. 


155 


1883: Gallinas Canyon, Sangre de Cristo Mountains 


The butterfly collections of the 1883 expedition re- 
main a mystery. Snow (1885) stated that in “July and 
August, 1883, our collections were made in the same 
locality as 1882—the Gallinas cafion, near the Las Ve- 
gas Hot Springs. My assistants were Messrs. L. L. Dy- 
che, W. H. Brown, W. C. Stevens, and [my son] W. A. 
Snow.” Lists of Lepidoptera and Coleoptera published 
for this trip included only those species that were not 
reported from 1882 (Snow 1885). Apparently no but- 
terflies satisfied this criterion because none were listed 
among the many beetles and moths. Similarly, the 
SEM contains no butterfly specimens with labels 
traceable to this expedition. Snow’s decision to neither 
report duplicates nor retain them in the collection 
would have been consistent with his normal practice. 


1884: Walnut Canyon and Elk Mountain 


The 1884 expedition included the same people as in 
1883, but this time they split into two groups and pro- 
duced butterflies from two geographically and ecolog- 
ically distinct localities. Dyche and Brown detrained at 
Las Vegas and returned to the site of their successful 
1882 and 1883 campaigns. Prof. Snow, his son Willie 
and W. C. Stevens traveled to Silver City in the south- 
west corner of New Mexico (Fig. 1). As far as we have 
been able to ascertain, and in contrast to previous ex- 
peditions, the 1884 butterfly specimens are not labeled 
to indicate month, year and locality of capture. Per- 
haps material from the two parties became mixed at 
some point, placing in doubt the provenance of indi- 
vidual specimens. Because Snow returned with butter- 
flies unique to southwestern New Mexico and re- 
ported them (Snow 1885), the authors were able to 
propose below a consistent labeling scheme for the 
1884 specimens. 

Little Walnut Creek. Snow “encamped about 
twelve miles north of Silver City, on the Walnut Creek, 
some three miles west of the divide which separates 
the Atlantic and Pacific slopes” (Snow 1885:65) (Fig. 
1). This coincides today with the Gila National Forest's 
Little Walnut Creek Picnic Ground, at about 6500 feet 
elevation in the Pinos Altos Mountains. 

Snow’s entire report for 1884 listed just 13 species 
of butterflies, all from Little Walnut Creek (Table 4a). 
His list was short considering that the area supports 
well over 100 species (e.g., Hubbard 1965, Ferris 
1976, Zimmerman 2001) and that summer rains usu- 
ally make August a productive month. Snow (1885:69) 
explained that “the season was a very dry one, there 
being hardly enough rain, except on one occasion, to 
thoroughly wet the canvas of our tent in the Walnut 


156 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 3. Butterflies collected during Snow’s 1882 expedition to Gallinas Canyon, San Miguel County, NM. 


Reported by Number of 
Taxon Snow (1883) as SEM! specimens Importance 
Thirybes pylades (Scudder, 1870) Eudamus Pylades Scud. 1? Ist NM specimen 
Erynnis horatius (Scudder & Burgess, 1870) not reported i? lst NM specimen 
Erynnis pacuvius Thanaos juvenalis E+ 
Thanaos Pacuvinus Lintn. 

Pyrgus communis (Grote, 1872) Pyrgus tesselatta Scud. Ist NM report 
Oarisma edwardsii (Barnes, 1897) not reported lee lst NM specimen 
Oarisma garita (Reakirt, 1866) Thymelicus Hylax Edw. I 1st NM specimen 
Hesperis comma (Linnaeus, 1758) Pamphila Comma var. Juba Scud. ° 
Hesperia viridis (W. H. Edwards, 1883) Pamphila Comma n. var. 
Polites themistocles (Latreille, 1824) Pamphila Cernes Bd.-Lec. 1 1st NM specimen 
Poanes taxiles (W. H. Edwards, 1881) Pamphila Taxiles Edw. ‘ 2 Ist NM specimens 
Paratrytone snowi (W. H. Edwards, 1877) Pamphila Snowi Edw. Ist NM report 
Euphyes vestris (Boisduval, 1852) Pamphila Metacomet Harr.* 1st NM report 
Amblyscirtes aenus W. H. Edwards, 1878 Amblyscirtes aeneus Edw. 1st NM report 
Amblyscirtes cassus W. H. Edwards, 1883 not reported 1 1st NM specimen 
Amblyscirtes phylace (W. H. Edwards, 1878) Pamphila Phylace Edw. 2 lst NM specimens 
Pterourus multicaudatus (W. F. Kirby, 1884) Papilio Daunus Edw. 1 
Pterourus rutulus (Lucas, 1852) Papilio Rutulus Bd. 1 
Neophasia menapia (C. and R. Felder, 1859) Pieris Menapia Feld. 1st NM report 
Pontia sisymbrii (Boisduval, 1852) Pieris Sisymbri Bd. 1 Ist NM specimen 
Pontia occidentalis (Reakirt, 1866) Pieris occidentalis Reak. lst NM report 
Pieris napi (Linnaeus, 1758) Pieris oleracea Bd.° 1 Ist NM specimen 
Anthocharis thoosa (Scudder, 1878) Anthocaris Thoosa Scud. 1 1st NM specimen 
Euchloe ausonides (Lucas, 1852) Anthocaris Ausonides Bd. 1 Ist NM specimen 
Colias eurytheme Boisduval, 1852 Colias Eurytheme Bd. 1st NM report 
Zerene cesonia (Stoll, 1790) Colias Caesonia Stoll. 
Nathalis iole Boisduval, 1836 Nathalis Iole Bd. 
Lycaena arota (Boisduval, 1852) Chrysophanus Ianthe Edw. 1 Ist NM specimen 
Lycaena helloides (Boisduval, 1852) Chrysophanus helloides Bd. 1 lst NM specimen 
Hypaurotis crysalus (W. H. Edwards, 1873) Thecla Crysalus Edw. 
Satyrium calanus (Hiibner, 1809) Thecla Calanus Hubn. 2 1st NM specimens 
Satyrium behrii (W. H. Edwards, 1870) Thecla Behrii Edw. 1st NM report 
Callophrys affinis (W. H. Edwards, 1862) Thecla Apama Edw. 1 lst NM specimen 
Mitoura grynea (Hiibner, 1819) 

ssp. siva (W. H. Edwards, 1874) Thecla Siva Edw. 1 
Incisalia eryphon (Boisduval, 1852) Thecla Eryphon Bd. 1 Ist NM specimen 
Leptotes marina (Reakirt, 1868) Lycaena marina Reak. Ist NM report 
Everes amyntula (Boisduval, 1852) Lycaena Amyntula Bd. 
Glaucopsyche lygdamus (Doubleday, 1841) 

ssp. oro (Scudder, 1876) Lycaena Lygdamus var. Oro Scud. 1st NM report 
Lycaeides melissa (W. H. Edwards, 1873) Lycaena Melissa Edw. 1st NM report 
Icaricia lupini (Boisduval, 1869) Lycaena Acmon West-Hew. 
Agriades franklinti (Curtis, 1835) Lycaena orbitulus Von Pr. 1 Ist NM specimen 
Apodemia nais (W. H. Edwards, 1876) Nemeobius Nais Edw. il 
Euptoieta claudia (Cramer, 1775) Euptoieta Claudia Cram. 1 
Speyeria aphrodite (Fabricius, 1787) 

ssp. ethne (Hemming, 1933) Argynnis Alcestis Edw. 1 Ist NM specimen 
Speyeria hesperis (W. H. Edwards, 1864) 

ssp. electa (W. H. Edwards, 1878) Argynnis Electa Edw. 
Speyeria mormonia (Boisduval, 1869) 

ssp. eurynome (W. H. Edwards, 1872) Argynnis Eurynome Edw. 1 lst NM specimen 
Poladryas minuta (W. H. Edwards, 1861) Melitaea minuta Edw. 1st NM report 
Thessalia fulvia (W. H. Edwards, 1879) Melitaea Fulvia Edw. 1 lst NM specimen 
Chlosyne nycteis (Doubleday and Hewitson, 1847) Phyciodes Nycteis var. 1st NM report 
Phyciodes cocyta (Cramer, 1777) Phyciodes Tharos n. var.'° 1st NM report 
Phyciodes pratensis (Behr, 1863) 

ssp. camillus W. H. Edwards, 1871 Phyciodes Camillus Edw. 1st NM report 
Euphydryas anicia (Doubleday and Hewitson 1848) Melitaea Nubigena Behr 1st NM report 


Polygonia gracilis (Grote and Robinson, 1867) 
ssp. zephyrus (W. H. Edwards, 1870) Grapta Zephyrus Edw. 


VOLUME 56, NUMBER 3 


Taxon 


Nymphalis milberti (Godart, 1819) 
Nymphalis antiopa (Linnaeus, 1758) 
Nymphalis californica (Boisduval, 1852) 
Vanessa virginiensis (Drury, 1773) 
Vanessa cardui (Linnaeus, 1758) 
Limeitis weidemeyerii (W. H. Edwards, 1861) 
Cyllopsis pertepida (Dyar, 1912) 
Coenonympha ochracea W. H. Edwards 
Cercyonis pegala (Fabricius, 1775) 
Cercyonis oetus (Boisduval, 1869) 

ssp. charon (W. H. Edwards, 1872) 
Erebia epipsodea Butler, 1868 
Neominois ridingsii (W. H. Edwards, 1865) 
Oeneis chryxus (Doubleday and Hewitson, 1849) 
Danaus plexippus (Linnaeus, 1758) 
Danaus gilippus (Cramer, 1775) 


TABLE 3. Continued. 


Number of 
SEM! specimens 


Reported by 
Snow (1883) as 


Vanessa Milberti Godt. 

Vanessa Antiopa L. 

Vanessa Californica Bd. 1 
Pyrameis Huntera Drury 

Pyrameis cardui L. 

Limenitis Weidemeyerii Edw. 

Neonympha Henshawii Edw. 

Coenonympha ochracea Edw. 

Satyrus Nephele Kirby 


Satyrus Charon Edw. 
Erebia epipsodea Butler 
Hipparchia Ridingsii Edw. 
Chionobas Uhleri Reak. 
Danais Archippus Fab. 
Danais Berenice Cram. 


157 


Importance 


1st NM report 
Ist NM specimen 


Ist NM report 
Ist NM report 


Ist NM report 
lst NM report 
Ist NM report 
Ist NM report 


"SEM = Snow Entomological Museum at the University of Kansas. 


* Snow also reported “Eudamus Bathyllus Sm.-Abb.” This specimen of T. pylades has an old label misidentitying it as T. bathyllus. 
° An old label misidentifies this specimen as E. martialis (Scudder), which Snow (1883) did not report. 
+ E. juvenalis does not occur in New Mexico; without a specimen, there is no way to know to what Snow referred. 


° An old label misidentifies this specimen as O. garita. 


° Although no specimen was found, this was likely a routine H. comma rather than a range-extending H. juba (Scudder). 
*“ Snow also reported “Poanes Zabulon Bd.-Lec.” which does not occur in New Mexico. 

° Snow also reported “Pamphila bimacula Gr-Rob.” which does not occur in New Mexico. 

° Snow also reported “Pieris Napi var. pallida Scud.,” which was synonymized. 

10 Snow also reported “Phyciodes tharos var. Marcia Edw.,” which refers to the same species. 


creek cafion during a stay of five weeks. The bed of the 
creek was dry for long distances.” Eight of the 13 
species reported by Snow (1885) were found in the 
SEM collection, each pinned to a label tersely typeset 
in black ink stating merely “New Mexico. F. H. Snow.” 

In 2002, the SEM had 15 additional specimens with 
“New Mexico. F. H. Snow.” labels, which Snow (1885) 
did not report. Habitat preferences and geographic 
distributions (e.g., Cary & Holland 1994, Toliver et al. 
1994) suggest that Little Walnut Creek was the only 
one of Snow's New Mexico destinations capable of 
producing several species bearing the “New Mexico” 
labels (Table 4b). These additional 11 species bring the 
1884 Little Walnut Creek catch to 24 species. To that 
it seems reasonable to add a small series of Cyllopsis 
pertepida specimens in the AMNH with like “New 
Mexico” labels (Miller 1974:72). Together, these spec- 
imens support a common origin for the “New Mexico. 
F. H. Snow.” labels. 

Among the final tally for Snow's Little Walnut Creek 
party were three new species: Atrytonopsis lunus (W. 
H. Edwards), Piruna polingii (Barnes) and Erynnis 
meridianus Bell, which were described in 1884, 1900 
and 1927, respectively (Miller & Brown 1981). The 
biggest surprise was Eurema salome (C. Felder & R. 
Felder), a subtropical stray for which Snow’s specimen 


remains the only New Mexico record. This specimen 
was attributed to New Mexico by Ehrlich and Ehrlich 
(1961:64) but without substantiating data, leaving re- 
cent workers skeptical (e.g., Scott 1986:207, Toliver et 
al. 1994:203). The specimen in question was in the 
SEM in 2002 pinned to a “New Mexico. F. H. Snow.” 
label and to a second label stating it was “illustrated in 
Ehrlich’s ‘How to Know the Butterflies’.” 

Elk Mountain. Gallinas Canyon was the starting 
point for the other section of the 1884 expedition, 
which apparently did no collecting there. Dyche ex- 
plained that his party “collected part of the summer at 
Harveys ranch, on top of “Baldy mountain, about 
twenty-five miles northwest of Las Vegas” (Dyche 
1909:43). The Harvey property is at 9500 feet eleva- 
tion near the head of Cascade Canyon, a tributary to 
Gallinas Creek (U.S. Forest Service 1994). From 
there, Elk Mountain was a brief six-mile horseback 
ride to the west. At 11,700 feet elevation, Elk Moun- 
tain is the nearest peak sufficiently treeless to qualify 
as “bald.” Dyche and Brown traveled on, spending 
“about six weeks of our time thirty to forty miles to the 
northwest of Harvey's ranch, on the head waters of the 
Pecos river” (Dyche 1909:43). There is no suggestion 
that they reached the highest peaks, which exceed 
13,000 feet elevation. 


158 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


TABLE 4A. Butterflies collected during Snow’s 1884 expedition to Little Walnut Creek as reported by Snow (1885). 


Taxon 


Autochton cellus (Boisduval and Leconte, 1837) 
Cogia caicus (Herrich-Schiffer, 1869) 

ssp. moschus (W. H. Edwards, 1882) 
Erynnis pacuvius (Lintner, 1876) 
Piruna polingii (Barnes, 1900) 
Amblyscirtes aenus W. H. Edwards, 1878 
Atrytonopsis lunus (W. H. Edwards, 1884) 
Battus philenor (Linnaeus, 1771) 
Atlides halesus (Cramer, 1777) 
Erora quaderna (Hewitson, 1868) 
Leptotes marina (Reakirt, 1868) 
Hemiargus isola (Reakirt, 1866) 
Celastrina ladon (Cramer, 1780) 


Reported by 
Snow (1885) as 


Eudamus cellus Bd.-Lec. 


Eudamus moschus Edw. 


Thanaos pacuvius Lintn. 


Pholisora, n. sp. 


Amblyscirtes aenus Edw. 


Pamphila lunus Edw. 
Papilio philenor L. 
Thecla halesus Cram. 
Thecla laeta Edw. 
Lycaena marina Edw. 
Lycaena alce Edw. 
Lycaena neglecta Edw. 


Number of 
SEM! specimens 


pp pe 


Importance 
1st NM report 
Ist NM specimen 
Ist NM specimen 
lst NM specimen 
Ist NM specimen 
Ist NM report 


Ist NM specimen - 


Ist NM specimen 
Ist NM specimen 


Limenitis arthemis (Drury, 1773) 
ssp. arizonensis (W. H. Edwards, 1882) 


' SEM = Snow Entomological Museum at the University of Kansas. 


The expedition report (Snow 1885) listed no butter- 
flies collected by this party, but we believe a few but- 
terflies captured by Dyche and Brown reside in the 
SEM collection, linked to the 1884 expedition by 
vague “New Mexico. F. H. Snow.” labels (Table 4c). 
Their habitat requirements associate them with the 
Canadian Zone west of Las Vegas, rather than the Up- 
per Sonoran and Transition Zone habitats of Little 
Walnut Creek. Some of these species also were caught 
by the 1882 expedition (Table 3). 

The “New Mexico. F. H. Snow.” labels may always 
be somewhat problematic. One could argue that cer- 
tain individual specimens with that label match up 
with published reports from 1881 or 1882. But it 
seems unlikely that the careful Snow would attach 
vague labels to one specimen in lieu of the informative 
labels pinned to the other specimens from that expedi- 
tion. Seeking parsimony for the “New Mexico. F. H. 
Snow.” labels, we suggest that they were printed for 
and affixed to one lot of specimens. 1884 was the only 
year that could have produced all those specimens and 
no specimens from 1884 are otherwise labeled. 

What might have prompted Snow to attach such 
general labels to this lot? Snow was a notoriously hard 
worker who accepted all challenges, leaving less time 
for his collections (Dyche 1909:43). In the early 1880s 
Snow taught botany, zoology, geology, natural history, 
meteorology and comparative anatomy—a burden of 
classes “which ought to have been taught by four men” 
(Hyder 1953:143). He also helped Kansas farmers deal 
with insect pests. Snow led no expeditions from 1885 
to 1889, but continued adding teaching duties until he 
was physically and mentally exhausted (Hyder 
1953:143). If Snow’s 1884 expedition concluded as 


Limenitis ursula Fab. var. Arizonensis Edw. 


1st NM report 


such expeditions became low priority, there may have 
been adverse consequences for specimen manage- 
ment and labeling. 

One remaining puzzle is Poanes zabulon, which 
Snow (1883) reported from his 1882 expedition (see 


TABLE 4B. Butterflies attributable to Snow's 1884 expedition to 
Little Walnut Creek, but not reported by Snow (1885). 


Number of 


Taxon SEM specimens Importance 


Erynnis meridianus 


Bell, 1927 1 Ist NM specimen 
Erynnis afranius 

(Lintner, 1878) 1 
Poanes taxiles 

(W. H. Edwards, 1881) 1 
Euphyes vestris 

(Boisduval, 1852) 1 
Amblyscirtes nereus 

(W. H. Edwards, 1876) 1 1st NM specimen 
Eurema mexicana 

(Boisduval, 1836) 1 
Eurema salome 

(C. and R. Felder, 1861) 1 Ist NM specimen 
Eurema nicippe 

(Cramer, 1779) iL 


Plebejus lupini 
(Boisduval, 1869) 
Thessalia theona 
(Ménétriés, 1855) 
ssp. thekla 
(W. H. Edwards, 1870) il 
Danaus gilippus 
(Cramer, 1775) 1 


bo 


Ist NM specimens 


Ist NM specimen 


lst NM specimen 


1 SEM = Snow Entomological Museum at the University of 
Kansas. 


VOLUME 56, NUMBER 3 


TABLE 4c. Butterflies collected by the Dyche party of Snow’s 
1884 expedition, but not reported by Snow (1885). 


Number of 

Taxon SEM! specimens Importance 
Polites origenes 

(Fabricius, 1793) yy) Ist NM specimens 
Colias scudderi 

(Reakirt, 1865) 1 lst NM specimen 
Erebia epipsodea 

(Butler, 1868) ) 
Oeneis chryxus 

(Doubleday and 1 

Hewitson, 1849) 
Oeneis uhleri 

(Reakirt, 1866) I 1st NM specimen 


' SEM = Snow Entomological Museum at the University of 
Kansas. 


Table 3). In 1978 Toliver located a specimen in the 
SEM and found it correctly determined, but its “New 
Mexico. F. H. Snow.” label links it to the 1884 expedi- 
tion. More worrisome, however, is that P. zabulon does 
not occur west of the Great Plains (e.g., Ferris & 
Brown 1980:104, Scott 1986:452). Poanes zabulon 
breeds in the vicinity of KU and the SEM has many 
specimens of local provenance. Snows New Mexico 
specimen likely is mislabeled and the report in error; 
this may be the exception that shows what a careful 
scientist Snow was. 


1894: Hop Canyon, Magdalena Mountains 


After a 10-year absence from New Mexico, Snow re- 
turned to the Magdalena Mountains in early August 
1894 with his son Frank, Ermine Case, Hugo Kahl and 
an unnamed student (Hyder 1953:280). They made 
camp in Hop Canyon, which is on the northwest flank 
of the Magdalena Mountains and descends through 
the town of Magdalena (Fig. 1). Snows camp may 
have been near the then-active lead mining camp of 
Kelly, about 30 air miles southwest of Socorro and 10 
air miles west of his Water Canyon camp of 1881, and 
the terminus of the local railroad at that time. 

Compared to that abbreviated effort 13 years ear- 
lier, this expedition went more smoothly. A fragment 
of a letter from Hop Canyon, dated 3 August 1894, 
was transcribed by one of Snow’s daughters many 
years later: “I think I was never so completely cut off 
from the world as here in this beautiful canyon . . . We 
have had no interruption of our successful entomolog- 
ical campaign except on Sunday and today. The latter 
interruption was in consequence Or A great storm 
which came upon us last night . . .. The dry bed of the 
canyon became an immense torrent in five minutes’ 
time and we were somewhat afraid that the flood 


159 


TABLE 5. Butterflies collected during Snow's 1894 expedition to 
Hop Canyon, Magdalena Mountains, Socorro County, New Mexico. 


Number of 


Taxon SEM! specimens Importance 
Erynnis pacuvius 
(Lintner) 1 
Celastrina ladon 
(Cramer, 1780) 2 
Lycaeides melissa 
(W. H. Edwards, 1873) ] 


Apodemia mormo 
(C. and R. Felder, 1859) 
ssp. mejicana 


(Behr, 1865) it Ist NM specimen 
Speyeria hesperis 

(W. H. Edwards, 1864) 

ssp. dorothea Moeck, 1947 1 1st NM specimen 


Polygonia gracilis 
(Grote and Robinson, 1867) 
ssp. zephyrus 


(W. H. Edwards, 1870) J 
Nymphalis milberti 

(Godart, 1819) 2 
Vanessa virginiensis 

(Drury, 1773) 1 


' SEM = Snow Entomological Museum at the University of 
Kansas. 


would reach our tents, but we were high enough up to 
avoid that catastrophe . . . I have never had so efficient 
a team for collection of insects. Will and Mr. Kahl take 
first place and the rest of us are not far behind. We 
have averaged about 500 pinned insects per day . . .” 

This expedition occurred while Snow was preoccu- 
pied as UK Chancellor and did not publish expedition 
results. Later, Snow (1907) published a comprehensive 
list of the Coleoptera collected on all six New Mexico 
expeditions including this one, but no 1894 Lepi- 
doptera have been published previously. 

The SEM contains nine examples of eight species 
from this expedition (Table 5). Most specimens have a 
typeset label stating “Magdalena Mts. N. M. Aug. °94. 
Snow.” Others are labeled with “Magdalena, New 
Mexico.” Neither label specifies elevation, but Mag- 
dalena is at 6600 feet and Kelly is at 7400 feet. Snow's 
practice in previous years suggests that locality labels 
are best interpreted as general descriptions of the area 
collected. Habitat associations suggest that Snow ven- 
tured to about 8000 feet elevation. 

A few 1894 specimens are known from other insti- 
tutional collections. A single specimen of Mitoura 
grynea siva (W. H. Edwards) was found by the senior 
author during review of collections at the NMNH ca. 
1985. The name of Snow's expedition assistant, H. 
Kahl, appears on labels of two Carnegie Museum 


160 


(CM) specimens of Mestra amymone (Ménétriés) 
from the “Magdalena Mts. N. M. Aug. 94. Snow.” 
(Toliver et al. 1994:387). 


DISCUSSION 


Westward extension of the railroad opened the door 
for Snows New Mexico expeditions. Snow was among 
the first entomologists to take advantage of 1000 miles 
of new track that penetrated and traversed New Mex- 
ico between 1879 and 1881 (Myrick 1990). With free 
passes donated by the Atchison, Topeka and Santa Fe 
Railroad (Hyder 1953:153), Snow probed the limits of 
railroad transportation in New Mexico in 1880, 1881, 
1884 and 1894. 

At the forefront of butterfly discovery in the Ameri- 
can Southwest, Snow succeeded in finding butterflies 
new to science. Lycaena cupreus snowi (W. H. Ed- 
wards) and Paratrytone snowi (W. H. Edwards) were 
named in his honor after he discovered them in Col- 
orado in the late 1870s (Snow 1879, 188la). He col- 
lected examples of several species in New Mexico be- 
fore they were formally described but, despite sending 
specimens to prominent scientists, most of Snow’s 
New Mexico material was not used to describe new 
species. Only Hesperia viridis was described based on 
specimens collected by Snow in New Mexico (Edwards 
1883). He also collected several taxa concurrent with, 
or shortly after, publication of formal descriptions. 

Most of Snow's extant New Mexico butterfly speci- 
mens were located in the SEM, whose modest hold- 
ings of North American butterflies made the search for 
80 Snow specimens reasonably profitable. In contrast, 
the few isolated specimens known from other institu- 
tional collections were encountered largely by acci- 
dent. Collections at the NUNH, CM and AMNH are 
much larger and the numbers of Snow butterflies 
much smaller. The NMNH has records of receiving 
many of Snow's Orthoptera, but no such records for 
butterflies (Marc Epstein, pers. com., February 2002). 

Workers in beetles and moths may find treasures 
among Snow’ material. In his New Mexico expedition 
reports, Snow's lists of beetles and moths dwarfed 
those of butterflies. His role in the discovery and de- 
scription of Daritis howardi (Hy. Edwards) (Arctiidae: 
Pericopinae) was documented elsewhere (Cary 2002). 
Snow collected beetles and moths at more places than 
he did butterflies, including Albuquerque and Socorro 
(Snow 1883, 1907). Rather than sit idle during evening 
stopovers in those towns, it would have been charac- 
teristic of Snow to find city lights or build large fires 
around which to collect beetles and moths all night. 
Some butterfly specimens eventually may turn up 
from these locations. 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Snow’s collections were the first substantive effort to 
document New Mexico butterflies. He recorded ap- 
proximately 90 species—still the largest single contri- 
bution to knowledge of the New Mexico fauna. Some 
of these probably were collected in New Mexico pre- 
viously, but most earlier reports lack meaningful docu- 
mentation and, except for a few types, are anecdotal 
and lack scientific value. Snow set a new standard by 
publishing his results and describing collectors, identi- 
fiers, locations and dates. He left dozens of specimens 
pinned with labels containing information still useful 
today. Subsequent New Mexico butterfly study would 
be based on this foundation. 


ACKNOWLEDGEMENTS 


The authors thank the staff of the Snow Entomological Museum 
at the University of Kansas for their generous assistance and hospi- 
tality. Staff of the Univ. ersity of Kansas Archives provided helpful in- 
formation, including several important letters. James Prewitt gra- 
ciously prepared the map in Fig. 1. Suggestions from two 
anonymous reviewers helped to make this paper more concise and 
complete. 


LITERATURE CITED 


Confusion persists regarding publication dates for the Transac- 
tions of the Kansas Academy of Science (TKAS). They were pub- 
lished every two years, so that manuscripts submitted in 1877 and 
1878 were published i in 1878 as Volume VI. One partial bibliography 
(Marvin 1909) confused the matter, giving dates for both even and 
odd years. Hyder (1953:269-273) dodged therssue by indicating the 
TKAS volume, but not the year of publication. We tried to give cor- 
rect dates below, but in case of confusion the reader should put 
more faith in the volume enumerated than the year of publication. 


Cary, S. J. 2002. Winslow J. Howard, pioneer New Mexico natu- 
ralist. J. Lepid. Soc. 56:49-52. 

Cary, S. J. & R. HOLLAND. [1994] 1992. New Mexico butterflies: 
checklist, distribution and conservation. J. Res. Lepid. 
31(1-2):57-82. 

DycuE, L. L. 1909. Dr. Snow as a collector, and his collections. 
Pages 39-45 in Memorial Addresses. TKAS XXII. 

Epwarps, W. H. 1883. Notes on Dr. Speyer’s paper. Canadian Ent. 
15:147-151. 

EHRLICH, P. R. & A. H. EHRLICH. 1961. How to know the butter- 
flies. W. C. Brown Co., Dubuque. 

FAUNTLEROY, G. 1999. Paving Canyon Road was an emotional tug 
of war. The Santa Fe Now Mexican. June 6, 1999. 

Ferris, C. D. 1976. A checklist of the butterflies of Grant County, 
New Mexico and vicinity. J. Lepid. Soc. 30(1):38-49. 

Ferris, C. D. & F. M. Brown. 1980. Butterflies of the Rocky 
Mountain States. U. of Oklahoma Press, Norman. 442 pp. 
HUBBARD, J. P. 1965. Some butterflies of the Pinos Altos Moun- 

tains, New Mexico. J. Lepid. Soc. 19(4):231-232. 

Hyper, C. K. 1953. Snow of Kansas: the life of Francis Huntington 
Snow with extracts from his journals and letters. U. of Kansas 
Press, Lawrence. 296 pp. 

MARVIN, F. O. 1909. The scientific papers of Dr. Francis Hunting- 
ton Snow. Pages 28-34 in Memorial Addresses. TKAS XXII. 

MILLER, E. 1909. Francis Huntington Snow, scientist. Pages 19-28 
in Memorial Addresses. TKAS XXII. 

MILLER, L. D. 1974. Revision of the Euptychiini (Satyridae) 2. Cyl- 
lopsis R. Felder, Bull. Allyn Mus. No. 20. 98 pp. 

MILLER, L. D. & F. M. Brown. 1981. A catalogue/checklist of the 
butterflies of America north of Mexico. The Lepidopterists’ So- 
ciety. Memoir No. 2. 280 pp. 


VOLUME 56, NUMBER 3 


Myrick, D. F. 1990. New Mexico’s railroads. Rev. ed. U of New 
Mexico Press, Albuquerque. 276 pp. 

Scort, J. A. 1986. Butterflies of North America, a natural history and 
field guide. Stanford Univ. Press, Stanford, Califormia. 583 pp. 

Snow, F. H. 1879. List of Lepidoptera collected in Colorado in 
June, July and August, 1876, by the Kansas University Scientific 
Expedition. TKAS VI:70-75. 

. 1881a. List of Lepidoptera, collected near Idaho Springs, 

Colorado, by the Kansas University Scientific Expedition of 

1879. TKAS VII:61-63. 

. 1881b. List of Coleoptera collected in Santa Fe Canon, 

New Mexico, by the Kansas University Expedition for 1880. 

TKAS VII:67-71. 

. 1883. Lists of Lepidoptera and Coleoptera, collected in 

New Mexico by the Kansas University Scientific Expeditions of 

1881 and 1882. TKAS VIII:35—45. 

. 1885. Lists of Lepidoptera and Coleoptera collected in 

New Mexico by the Kansas University Scientific Expeditions of 

1883 and 1884. TKAS IX:65-69. 


161 


. 1907. List of Coleoptera collected in New Mexico by the 
Entomological Expeditions of the University of Kansas. TKAS 
XX(2):165-189. 

STEVENS, W. C. 1909. Professor Snow as a teacher. Pages 34—36 in 
Memorial Addresses. TKAS XXII. 

To.iver, M. E., R. HOLLAND & S. J. Cary. 1994. Distribution of 
butterflies in New Mexico (Lepidoptera: Hesperioidea and Pa- 
pilionoidea). 2nd ed. Published by R. Holland, Albuquerque. 
439 pp. + appendices. 

U.S. ForEST SERVICE. 1994. Pecos Wilderness, Carson and Santa 
Fe National Forests. Map with 80-foot contour interval and 
scale of 1 inch to 1 mile. 2nd printing. 

ZIMMERMAN, D.A. 2001. An inventory of the butterfly species on 
the Gila National Forest, southwestern New Mexico. Report to 
Gila Nat. For. 112 pp. 


Received for publication 7 May 2001; revised and accepted 10 
March 2002. 


Journal of the Lepidopterists’ Society 
56(3), 2002, 162-165 


DISTRIBUTION OF THE DIANA FRITILLARY, SPEYERIA DIANA (NYMPHALIDAE) IN ARKANSAS, 
WITH NOTES ON NECTAR PLANT AND HABITAT PREFERENCE 


MATTHEW D. MorAN! AND CHARLES D. BALDRIDGE 
Department of Biology, Hendrix College, 1600 Washington Ave., Conway, Arkansas 72032, USA 


ABSTRACT. Investigation of the distribution, preferred nectar plants, and habitat associations of Diana Fritillary, Speyeria diana, Cramer 
in Arkansas was undertaken. Arkansas populations form a disjunct group separate from larger populations of this species in the Appalachian 
Mountains. Researchers have suggested that S. diana has declined over much of its range, including the Ozark and Ouachita Mountains of 
Arkansas, so that only a few populations are currently known in this area. Previous surveys found this butterfly in only nine Arkansas counties. 
We observed populations of this butterfly in 14 counties, 11 which were new county records, In addition, we confirmed populations in two coun- 
ties where the butterfly had not been recorded in over 20 years. Observations made during this study combined with previous survey work in- 
dicate that this species is distributed throughout the Ozark and Ouachita mountains in Arkansas, occupying 22 counties. Individuals were found 
to occupy two types of habitat; prairie and wetland, which appeared to contain specific nectar plants that S. diana prefers. We suggest that the 
loss of these habitats and associated nectar plants has been the primary cause of the butterfly’s decline, but with proper management and pro- 
tection of these habitats, the species may be increasing. Therefore, S. diana does not appear to be in immediate risk of extirpation in Arkansas 


although monitoring of existing populations is warranted. 


Additional key words: survey, Ozark Mountain, Ouachita Mountain, prairie, wetlands, habitat loss. 


Several North American fritillary butterflies have 
become endangered in the last century, often because 
of habitat alteration (Hammond & McCorkle 1983, 
Hammond 1995). One species of concern has been 
the Diana Fritillary, Speyeria diana Cramer 
(Nymphalidae). The historical range of S. diana ex- 
tended from the Chesapeake Bay region, across the 
southern Appalachians, through Tennessee, Kentucky, 
and into northern Georgia, Alabama, and Mississippi. 
Possible disjunct populations existed in Arkansas and 
southern Missouri, ending at the deciduous forest/ 
prairie ecotone in eastern Oklahoma and Kansas (Ho- 
vanitz 1963, Carlton & Nobles 1996). Several authors 
have suggested a great decline throughout much of the 
range of S. diana (Clark 1951, Shull 1987, Howe 1975) 
so that currently, populations exist only in the Ap- 
palachian Mountains and the Interior Highlands of the 
Ozark Plateau and Ouachita Mountains covering 
Arkansas, Missouri, and Oklahoma (Carlton & Nobles 
1996). Survey work conducted by Carlton and Nobles 
in 1996 found S. diana at several Ozark and Ouachita 
localities covering nine counties. The authors sug- 
gested that the populations were small and isolated 
and therefore at high risk for extinction. 

Speyeria diana emerges in late spring, mating oc- 
curs in early summer, after which males disappear and 
presumably die. Females are seldom seen during the 
rest of the summer months, but become active again in 
mid-autumn to oviposit. Eggs are deposited on the 
ground, the larvae hatch and overwinter as first sta- 
dium larvae. In early Spring, the larvae become active 
again, feed on various species of violet (Viola spp. L.), 
and pupate by mid-spring. There is one generation per 
year (Howe 1975). 


' Corresponding author. 


To document further the range of S. diana in the 
western portion of its range, we conducted surveys 
throughout Arkansas, focusing on areas where the but- 
terflies were not observed in the Carlton and Nobles 
(1996) survey. We also searched areas that had histori- 
cal records that have not been confirmed in recent 
years. Observations on the behavior of each butterfly 
observed were made, primarily nectar plant prefer- 
ences to determine possible habitat requirements. 


MATERIALS AND METHODS 


During the summers of 1997-1999 we performed 
extensive surveys of Arkansas habitats for S. diana. A 
total of 23 counties were surveyed by the authors near 
the known range of the butterfly. Additional records 
were provided by The Nature Conservancy, the Na- 
tional Forest Service, and other scientists in Arkansas. 
For each butterfly observed, we recorded its sex, 
noted its behavior when sighted, and the associated 
habitat. All observations were performed during June, 
July, and early August when adult butterflies are ac- 
tive. We began by focussing on areas with known (al- 
though old) records of S. diana, and then surveyed 
surrounding counties that did not have published 
records of this species. 

Surveys were performed on public lands by walking 
trails and driving back roads. In particular, areas that 
contained significant concentrations of nectar plants 
were searched carefully for S. diana. In-areas of pri- 
vate land, we drove slowly along roadsides searching 
for possible nectar plants. We searched a variety of 
available habitats including mature forest, wetlands, 
prairies, and human-disturbed areas. A total of 375 
man-hours were spent searching in the field. 


VOLUME 56, NUMBER 3 


RESULTS 


Individuals of Speyeria diana were observed each 
year of the study (1997-1999) at numerous sites across 
Arkansas. Records from observers other than the au- 
thors were also added in 2000-2001. Butterflies were 
found in 14 different Arkansas counties, 11 of which 
represent new county records. Two of these counties, 
Conway and Faulkner, have not had sightings of this 
species in over 20 years. Below are the initial observa- 
tions from each county (ie., first time we observed 
specimens). Observations were made by the authors 
unless otherwise indicated. 

Conway Co.; Petit Jean St. Park, West end of Bai- 
ley Lake, 10 July 1998, one female nectaring on but- 
tonbush (Cephalanthus occidentalis L.); Faulkner 
Co.; Camp Robinson National Guard Base, along 
Cemetery Road, forested wetland, 7 July 1997, four 
males and one female nectaring on C. occidentalis; Pu- 
laski Co.; Camp Robinson National Guard Base, along 
Clinton Rd., south of Clifton Mountain, forested wet- 
land, , 7 July 1997, one male nectaring on C. occciden- 
talis: Yell Co.; Mt. Nebo St. Park, Fern Lake near 
Summit Park Trail, 10 July 1998, one male nectaring 
on C. occidentalis; Johnson Co.; Ozark Highlands 
Trail in Hurricane Creek Wilderness, open glade, 17 
July 1997, one male nectaring on Purple Coneflower 
(Echinacea purpurea Moench), P. Kilgore; Logan 
Co.; Mt Magazine, one-half kilometer west of Signal 
Hill summit, open glade, 25 June 1997, 5 males and 1 
female nectaring on E. purpurea, MDM and P. Kil- 
gore; Howard Co.; Stone Road Glade Natural Area, 
June 1998, 3 males nectaring on Pale Purple Cone- 
flower (Echinacea pallida Britton) and 1 female nec- 
taring on Compass Plant (Silphiwm laciniatum L.), 
Douglas Zollner; Hempstead Co.; Grandview Prairie 
Wildlife Management Area, numerous males and fe- 
males sighted during summer of 1998 and 1999, Dou- 
glas Zollner; Clark Co.; Terre Noire Natural Area, 
June 1997, 1 male nectaring on E. pallida, and July 
1999, 3 females nectaring on S. laciniatum, Douglas 
Zollner; Jefferson Co.; Pine Bluff Arensal, June 1999, 
2 males and | female nectaring on E. pallida, Douglas 
Zollner; Saline Co.; Dry Lost Creek Preserve, late 
May 1999, 3 males nectaring on Arkansas calamint (Sa- 
tureia arkansana Nutt.), and June 1999, 1 female nec- 
taring on slender mountain mint (Pycnanthemum 
albescens Torr.), Douglas Zollner; Polk Co.; Ouachita 
National Forest, Forest Service Rd 1401, about 10 
miles south of Mena, AR, 1 July 2000, 1 female, Craig 
Rudolph; Garland Co.; Ouachita National Forest, 
Mazarn Creek and Forest Service Rd. 829, 2 June 
2000, 2 males, Craig Rudolph; Pike Co.; Ouachita Na- 


163 


Fic. 1. Range map of the Diana Fritillary (Speyeria diana) in 
Arkansas based on surveys from this study and previous records. Di- 
amonds = old observations from published literature; open circles = 
Carlton and Nobles (1996) observations, Closed circles = observa- 
tions from this study. 


tional Forest, Highway 84, 4.2 miles west of Salem, 
AR, 7 June 2001, 1 male, Craig Rudolph; Benton Co.; 
Wedington Natural Area, 24 July 2002, 1 male, Lori 
Spencer. These records indicate populations exist 
throughout much of the western one-half of Arkansas, 
primarily in the mountainous and foothill regions of 
the state. (Fig. 1). Our surveys in the eastern portion of 
the state failed to record any individuals except for the 
Jefferson County record provided by the Nature Con- 
servancy. 

Populations of S. diana were found in two types of 
habitats, prairie and wetland. In southwest Arkansas, 
many butterflies were found in prairie habitat. Individ- 
uals in four sites (Stone Road Glade, Grandview 
Prairie, Terre Noire, and Dry Lost Creek) were found 
in the year after prescribed burns. In the Ozark and 
Ouachita mountains, S. diana was associated with 
small natural prairie openings (e.g., Magazine Moun- 
tain, Hurricane Creek Wilderness) while those in cen- 
tral Arkansas were found in wetland areas (e.g., 
swamps in Camp Robinson, Mt. Nebo, and along Baily 
Lake). 

During our field observations we also recorded the 
activity of the butterflies. The vast majority of individ- 
uals were nectaring, however, butterflies were often 
observed on only a few species, with Buttonbush (C. 
occidentalis) and coneflowers (Echinacea spp.) the 
most commonly utilized plants (Table 1). Females 


164 


TABLE 1. Percentage of individual male and female S. diana ob- 
served on various nectar plants. N = number of individuals observed. 


Percent Males Percent Females 


Nectar Plant (N = 46) (N = 23) 
Cephalanthus occidentalis 56.5 26.1 
Echinacea purpurea 21.7 8.1 
Echinacea pallida 13.0 ie 
Pycnanthemum albescens 6.5 4.3 
Rubus sp. 2.2 0.0 
Silphium laciniatum 0.0 34.8 
Satureja arkansana 0.0 4.3 


were frequently observed on Compass Plant (S. 
laciniatum) on prairie sites. Several other plant species 
were present at our sites and used by other species of 
butterfly. Although there were more males than fe- 
males observed, this may not indicate an actual biased 
sex ratio as males have bright coloration and are there- 
fore easier to detect in the field. The habitat prefer- 
ence (wetland or prairie) seemed to be determined by 
the presence of preferred nectar plants, and not the 
habitat per se. 


DISCUSSION 


Our results indicate that the Diana Fritillary is more 
widespread than previously thought. Based on our 
newly constructed range map (Fig. 1) and habitat ob- 
servations, S. diana appears to range throughout the 
Ouachita and Ozark Mountains of Arkansas, where 
there is suitable wetland and/or prairie habitat with 
preferred nectar plants. We also performed extensive 
surveys throughout eastern Arkansas, but were unable 
to locate any individuals, even though preferred nectar 
plants utilized in other areas were present. 

It has been suggested that clearing of old-growth 
forest in the eastern United States has been primarily 
responsible for the decline of S. diana, due to larval 
host plant decline (Clark 1951, Howe 1975, Ham- 
mond & McCorkle 1983, Shull 1987). However, based 
on our observations, we find this explanation wanting. 
Speyeria diana larvae feed on several species of violets 
(Viola spp.), which are extremely common in 
Arkansas. Many of our observations and observations 
by Carlton and Nobles (1996) were in at least moder- 
ately disturbed habitat. The largest number of sight- 
ings in this study were in the Camp Robinson National 
Guard Base and surrounding areas, habitat that is 
mostly second growth forest and pasture, and is fre- 
quently burned ae to military firing exercises. Butter- 
flies were also observed after prescribed burning at 
several sites, further indicating that this type of distur- 
bance is not the primary cause of the butterfly’s de- 
cline. Reports from the National Forest Service sites 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


in western Arkansas indicate that butterflies become 
more common after prescribed burns are undertaken 
for Red Cockcaded Woodpecker habitat (C. Rudolph, 
pers. com.). We believe it is more likely that loss of 
wetland and prairie habitat and the associated loss of 
preferred nectar plants is the reason for this butterfly’s 
decline. Throughout the United States, large areas of 
wetland have been drained (Weller 1981, Tiner 1984) 
and most prairie habitat has been lost to farmland con- 
version or has undergone succession to forest because 
of fire suppression (Humphrey & Mehrhoff 1958, 
Bock & Bock 1995). 

It should be noted that we did not search for larvae, 
which may prefer quite different habitats compared to 
adults. The larval host plants (Viola spp.) are most com- 
mon in moist forest. In addition, many butterflies (es- 
pecially females) appear to prefer deep, shady forests 
during times of strong sunlight (P. C. Hammond, pers. 
com.). Therefore, high quality forest in the vicinity of 
quality nectar plants (i.e., mixture of forested and open 
habitats) may be important for this species. 

The Diana Fritillary has an unusually long adult life 
span, especially females, which are observed from 
June to October (up to 5 months). Many long-lived 
butterflies require high quality nectar sources (e.g., 
Heliconius, Gilbert 1972), and we suggest this is the 
case for S. diana. The loss of prairie and wetland habi- 
tats, and subsequent loss of nectar plants may have 
contributed to the decline of this species throughout 
much of its range. 

While our survey indicates S. diana is more wide- 
spread than previously thought, it is still a relatively 
rare butterfly. There are now 22 known populations in 
Arkansas, most of which are on public land. We sus- 
pect that further survey work will discover additional 
populations. It is unclear if the species’ population is 
increasing or has simply been overlooked in the past. 
The latter may be the case, as males resemble the 
Great Spangled Fritillary, Speyeria cybele Fabricius 
and females are rather secretive and resemble several 
other species, especially the Red-Spotted Purple, 
Basilarchia astyanax Drury. It is also possible that 
changes in habitat management are benefiting this 
species, as prescribed burning has become more com- 
mon and wetlands are better protected. While S. diana 
does not appear in immediate danger of extinction in 
the Ozark and Ouachita areas, future monitoring ef- 
forts will be required to determine if existing popula- 
tions are stable. 


ACKNOWLEDGMENTS 


Douglas Zollner (The Nature Conservancy), Craig Rudolph (U.S. 
Forest Service), and Lori Spencer provided some of the county 


VOLUME 56, NUMBER 3 


records for this study. Parker Kilgore and Laura Skelton enthusiasti- 
cally helped in field surveys. L. Spencer provided valuable criticisms 
to an earlier version of this manuscript. Paul C. Hammond and Jim 
Dunford provided helpful reviews. 


LITERATURE CITED 


Bock, J. H. & C, E. Bock. 1995. The challenges of grassland con- 
servation. In A. Joern and K. H. Keeler (eds.) The changing 
prairie: North American grasslands. Oxford University Press, 
New York, NY. 

Carton, C. E. & L.S. Noses. 1996. Distribution of Speyeria di- 
ana (Lepidoptera: Nymphalidae) in the highlands of Arkansas, 
Missouri, and Oklahoma, with comments on conservation. En- 
tomol. News 107:213-219. 

Ciark, A. H. 1951. The Butterflies of Virginia. Smithsonian Mis- 
cellaneous Collections 116:1—95. 

GILBERT, L. E. 1972. Pollen feeding and reproductive biology of 
Heliconius butterflies. Proc. Nat. Acad. Sci. 69:1403—1407. 


165 


HAMMOND, P. C. 1995. Conservation of biodiversity in native prairie 
communities in the United States. J. Kans. Entomol. Soc. 68:1-6. 

HAMMOND, P. C. & D. V. McCorkLeE. 1983. The decline and ex- 
tinction of Speyeria populations resulting from human environ- 
mental disturbances (Nymphalidae: Argynninae). J. Res. Lepid. 
29: I\ 7-294. 

Hovanitz, W. 1963. Geographical distribution and variation of the 
genus Argynnis diana. J. Res. Lepid. 1:201—208. 

Howe, W. H. 1975. The butterflies of North America. Doubleday, 
Garden City, NY. 

Humpueey, R. R. & L. A. MEHRHOFF. 1958. Vegetation changes on 
a southern Arizona grassland range. Ecology 39:720-726. 

SHULL, E. M. 1987. The butterflies of Indiana. Indiana University 
Press. Bloomington, IN. USA. 

TineER, R. W. 1984. Wetlands of the United States: current status 
and recent trends. U.S. Department of Interior, Fish and 
Wildlife Service, Washington, DC. 

WELLER, M. W. 1987. Freshwater marshes. University of Min- 
nesota Press, Minneapolis. 


GENERAL NOTES 


Journal of the Lepidopterists’ Society 
56(3), 2002, 166-170 


EVIDENCE FOR THE NEGATIVE EFFECTS OF BT 
(BACILLUS THURINGIENSIS VAR. KURSTAKI) ON A NON-TARGET 
BUTTERFLY COMMUNITY IN WESTERN OREGON, USA 


Additional key words: Bt, species richness, butterfly abundance, diversity. 


Bacillus thuringiensis var kurstaki (Bt) is a gram- 
positive bacterium commonly occurring in soil (Martin 
& Travers 1989) and plant leaves (Smith & Couche 
1991) with known pathogenic properties for butterfly 
and moth larvae. Generally, when Bacillus bacteria ex- 
perience starvation or stressful conditions they form a 
spore, an inactive resting stage (Brock 2000), that is 
lethal to most Lepidoptera larvae when ingested. Con- 
sumption of Bt spores by butterfly and moth larvae ac- 
tivates bacterial Cry proteins (toxins) by proteolysis 
from the larva’s gut enzymes (Tojo & Aizawa 1983). 
Once the bacteria’s toxins are activated by proteolysis, 
the proteins bind to the apical brush border gut mi- 
crovillae cells (Hofmann et al. 1988) and form an ion 
channel that disrupts solute concentrations within the 
cell. Ion pore size increases with increasing gut alka- 
linity resulting in decreasing solute control with lar ger 
pore sizes Gciware et al. 1993). The lack of eallteli 
solute control causes water to enter the cell and lyses 
cells having sufficient numbers of Bt ion pores. Pre- 
sumably, the loss or dysfunction of gut epithelial cells 
causes mortality of infected Lepidoptera species. 

B. thuringiensis var. kurstaki is commonly employed 
as a biological control agent for lepidopteran pests on 
agricultural plant species, but other Bt variants are 
pathogenic to mosquito and black flies (Hershey et al. 
1998), coleopterans, hymenopterans, and orthopterans 
(Schnepf et al. 1998). Bt is suspected of causing ab- 
normally high larval mortality rates in non-target but- 
terfly species like Monarchs, Danaus slices IL, 
(Danaidae) (Losey et al. 1999), various swallowtail 
species (Papilio) (Johnson et al. 1995, Peacock et al. 
1998), and a host of other butterfly and moth species 
(Wagner et al. 1996, Peacock et al. 1998, Whaley et al. 
1998). Miller (1990, 1992) observed a decrease in lar- 
val density, diversity, and species richness of lepi- 
dopteran g guilds feeding on Quercus garryana Douglas 
(Fagaceae) and Ceanothus velutinus Douglas (Rham- 
naceae) following aerial Bt application. Bio-engineer- 
ing of genes encoding the Bt Cry toxins into com 
plants and the dispersal of transgenic pollen into 
hedgerows bordering cornfields is linked to increased 
larval mortality of native butterfly species when pollen 


coated leaves are consumed by larvae (Losey et al. 
1999, Hansen Jesse & Obrycki 2000). 

Elevated larval death rates due to Bt not only im- 
pacts butterfly and moth populations, but is also capa- 
ble of altering higher trophic level interactions with in- 
sectivorous species. Sorex cinereus (masked shrew) 
males migrated out of a Bt treatment area and all 
shrews altered their prey selection from Lepidoptera 
larvae to less preferred hemipterans and hymenopter- 
ans after a Bt spray reduced larval abundance (Bellocq 
et al. 1992). Parent birds fed their nestlings fewer 
caterpillars in Bt treated areas (Gaddis & Corkran 
1986, Rodenhouse & Holmes 1992, Nagy & Smith 
1997) or increased their foraging times compared to 
birds nesting in untreated areas (Holmes 1998). Al- 
though a decrease in Lepidoptera larval abundance 
did not appear to effect the average nestling weight, 
survivorship through fledging, or egg size in birds from 
a Bt treated area (Gaddis & Corkran 1986, Roden- 
house & Holmes 1992, Nagy & Smith 1997), lower fat 
reserves were detected from birds in a Dimilin (a lep- 
idopteran larvicide with impacts similar to Bt) treated 
forest (Whitmore et al. 1993). 

Lepidoptera larvae, particularly early instars, experi- 
ence the acute effects of Bt toxins and therefore have 
received the bulk of the attention from researchers. 
Conversely, adult non-target Lepidoptera populations 
in Bt treated sites have received very little attention, 
possibly due to methodological problems accompany- 
ing the sampling of adult abundance over potentially 
large spray areas. However, Whaley et al. (1995) noted 
substantial reductions in adult population sizes for In- 
cisalia fotis Strecker (Lycaenidae) and Neominois rid- 
ingsti Edwards (Satyridae) i in Utah following a Bt spray. 

Increased larval mortality from Bt use should result 
in the reduction of adult non-target Lepidoptera 
populations. The butterfly community should respond 
to eradication dosages of Bt by displaying a reduction 
in species richness, diversity, and adult abundance in 
the years following the spray. Furthermore, because Bt 
will likely elevate arortalliy rates of all butterfly species 
in the community, reductions in species abundance 
should be synchronized among species. I monitored a 


VOLUME 56, NUMBER 3 


relatively small Bt spray, approximately 32 ha, that oc- 
curred in the spring of 1997 in western Oregon to as- 
sess the effects of Bt on the adult non-target butterfly 
community. 

Study site description. Schwarz Park, situated 
downstream of Dorena Reservoir, Lane Co. Oregon, is 
a 32.4 ha (80 acres) Bt treatment area for a local gypsy 
moth infestation. The Schwarz Park Bt spray site con- 
tains three different habitats that presumably received 
equal dosage treatments, a “groomed,” “prairie,” and 
“spillway” habitat. The groomed habitat consists of 
14.2 ha (35 acres) of lawn and Pseudotsuga menziesii 
(Mirbel) Franco (Pinaceae), Douglas-fir, forest main- 
tained for camping and recreational use. The spillway 
habitat is 8.1 ha (20 acres) of young riparian forest in- 
terspersed with native upland prairie and cliff face. An 
area of 10.1 ha (25 acres) of degraded upland prairie 
with a narrow swath (0.8 ha) of riparian habitat and 
young Douglas-fir forest compose the remaining 
prairie habitat. The Bt treatment area is bounded by 
reservoir water to the east, and sporadically clearcut 
Douglas-fir forests to the north, west, and south. Ex- 
cluding the nearby residential developments and an 
U.S. Forest Service tree genetics research site, Schwarz 
Park is the only area within 5 km that is suitable upland 
prairie butterfly habitat. On 30 April, 8 May, and 20 
May of 1997, three aerial applications of Foray 48B 
(Bt) at 24 B.I.U.s were administered by helicopter at 
least 15 m (50 feet) above the tree canopy over the 
Schwarz Park spray area (Bai & Johnson 1997). 

Adult surveys and analysis. The 32.4 ha spray 
area was subdivided into three habitats (spillway, 
prairie, and groomed) that were systematically sur- 
veyed for adult butterflies once every three to four 
weeks beginning in late April and ending in early June 
from 1997 through 2000. Target flight periods for sam- 
pling were late April, mid May, and late May/early 
June that correspond with the spring butterfly com- 
munity flight period. Temporal separation of adult 
sampling dates by approximately two weeks ensured 
that most adult butterflies would not be counted twice 
because the butterflies from the first survey period 
would likely be dead by the second survey period. In 
most instances, adult butterflies were accounted for 
and identified without capturing or disturbance. By 
avoiding injury and death of the adult population 
through surveyor disruption, butterfly reproductive ca- 
pability was not diminished through capturing or dis- 
turbance while ovipositing. Survey days were re- 
stricted to climate conditions that favored the flight of 
adult butterflies while maintaining the seasonal timing 
of the initial survey periods from the original spray 


167 


TABLE 1. Butterfly diversity for the Schwarz Park spray area. * = 
significant difference in diversity compared to the spray year, 1997. 


Year Shannon Year Comparison 
Diversity (H’) (modified t-test) of H’ 

1997 0.9840 

*1997 x 1998 t = 2.77 P <0.01 
1998 0.7408 

*1997 x 1999 t = 4.92 P <0.0001 
1999 0.5497 

1997 x 2000 t = 1.59 P > 0.05 

2000 0.8607 


year. Adult surveys were performed from 1030 h-1630 h 
and occurred on days when temperatures were near or 
above 21°C (70°F) under sunny conditions. Butterfly 
binomial nomenclature was synonymous with the lo- 
cally accepted names published by Hinchliff (1994). 

Adult surveys were conducted systematically by walk- 
ing transects approximately 5 m apart progressing from 
one end of a subarea, defined by physical barriers, to the 
other end. Subareas, nested inside the three habitats, 
were ().8—2.0 ha and delineated by physical boundaries 
so survey replication within and between sampling years 
was consistent. If large numbers of butterflies occupied 
a subarea (only occurred once in 1997), all butterflies 
within the site were captured in an aerial insect net 
and held until the butterflies in the subarea were ac- 
counted for, identified to species, and then released. 
Once all transects in a subarea were surveyed, butter- 
fly data gathering was finished for that particular sub- 
area and then began anew in the next subarea. 

The spray year, 1997, was used as baseline data for 
yearly pairwise comparisons because it was assumed 
that the majority of the adult spring butterfly species 
would have overwintered as a pupae or late instar lar- 
vae, and would thus be unaffected by the timing of the 
Bt spray. Butterfly abundance was estimated by divid- 
ing the total number of butterflies counted per survey 
by the spray area (32.4 ha). Butterfly diversity was cal- 
culated for each year using the Shannon Diversity In- 
dex (Magurran 1988). Yearly diversity measurements 
were compared for significant pairwise differences be- 
tween the spray year and the three postspray years us- 
ing a modified t-test described in Zar (1996). 

Community response. Diversity was significantly 
lower in the two postspray years, 1998 (P < 0.01) and 
1999 (P < 0.0001), when compared pairwise to the 
spray year 1997 (Table 1). The decline in diversity of 
the non-target butterfly community immediately fol- 
lowing the spray suggests that the Bt application nega- 
tively impacted the local butterfly community. Butter- 
fly species richness throughout the four-year study 


168 


period was also substantially lower in the two years fol- 
lowing the Bt spray (Appendix). In 1997 there were 22 
species detected in the spray area, but in the two fol- 
lowing postspray years species richness had dropped to 
only eight species (Appendix). In the year 2000, three 
years following the initial spray, species richness was 
19, a level similar to that measured in the spray year 
(Appendix). Furthermore, butterfly density was also 
substantially lower in the two postspray years com- 
pared to the spray year. In 1997 there were 5.49 indi- 
viduals/ha, while in 1998 and 1999, there were only 
1.27 and 2.68 individuals/ha respectively. However, 
there were more butterflies per hectare in 2000, 8.18 
individuals/ha, than there were in the spray year, sug- 
gesting that the assumption of a no treatment effect on 
the adult butterfly community during the spray year 
was violated with some species. 

Butterfly diversity, species richness, and density all 
showed similar reductions in numbers following the 
year following the Bt spray, but the lack of a suitable 
control area to statistically account for stochastic ef- 
fects on an untreated butterfly community is a draw- 
back to the this study. Despite a lack of a suitable con- 
trol area, fair numbers of Mitoura grynea (6 
individuals), Boloria epithore chermocki (8 individuals), 
Euphydryas chalcedona colon (18 individuals), Everes 
comyntas (10 individuals), Parnassius clodius claudi- 
anus (4 individuals), and Papilio rutulus (5 individu- 
als), were detected just outside the spray boundary in 
1998 during one sampling day. However, no individu- 
als of the same butterfly species were found within the 
Bt treatment area (Appendix), suggesting the detri- 
mental effects of Bt on the local butterfly populations. 

Further evidence for the effects of Bt on adult 
populations are demonstrated by comparing the yearly 
population size and lifehistory of the two dominant 
butterfly species, Ceononympha tullia eunomia and 
Glaucopsyche lygdamus columbia. Ceononympha t. 
eunomia was in the larval life stage during the Bt spray, 
so the small adult population in the spray year was 


likely the effect of the Bt treatment. The abundance of 


C. t. eunomia was lowest in the spray year and steadily 
increased with time following the Bt spray (Appendix). 
Comparing C. t. ewnomia to another dominant species, 
Glaucopsyche lygdamus columbia, that was in the pu- 
pal stage when the spray occurred (adults would not 
be affected in the spray year), a different trend in adult 
abundance occurred. Glaucopsyche |. columbia adult 
populations were high in the spray year and low the 
two following years ( Appendix), indicating that the Bt 
treatment did affect the competitively dominant 
species of the butterfly community and likely the com- 
petitively inferior species as well. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


The data from the Bt spray (Appendix) evince that 
an overall depression in butterfly abundance occurred 
in the years following the spray. Moreover, the de- 
creasing trend among the majority of the butterfly 
species was synchronized, implying that a large scale 
disturbance event affected the community. Outside of 
seasonality, synchronized patterns in species abun- 
dance are not expected to occur unless some cata- 
strophic ecological event affects the entire community. 
Butterfly community data from a five year study in 
Ecuador demonstrate the asynchronous nature of the 
component species in a butterfly community under 
natural conditions (DeVries & Walla 2001). Pollard 
(1984) collected butterfly species abundance data for 
numerous sites throughout England for seven years, a 
Temperate Zone with a butterfly community similar to 
western Oregon, and the butterfly species did not all 
follow the same pattern of relative abundance in- 
creases and declines between years. The only known 
factor that had the potential to cause a community 
wide concurrent decrease in species abundance at 
Schwarz Park was the Bt spray. 

Miller (1990, 1992), Peacock et al. (1998), and Wha- 
ley et al. (1998) predicted local species extinction of 
non-target Lepidoptera in response to repeated Bt ap- 
plication events. Locally distributed, monophagous, 
and rare Lepidoptera species are the ones most likely 
to experience a resultant Bt induced extinction be- 
cause they occupy narrow ecological niches. It appears 
that a localized extinction of three non-target butterfly 
species transpired within Schwarz Park following the 
spray. Mitoura grynea, Parnassius clodius claudianus, 
and Phyciodes p. pratensis were found in 1997, but 
were not detected inside Schwarz Park the three years 
following the spray (Appendix). Larvae of M. grynea 
use Calocedrus deccurens (Torr.) Florin. (Cupres- 
saceae), P. clodius claudianus uses Dicentra formossa 
(Andr.) Walp. (Papaveraceae), and P. pratensis uses 
Aster hallii (Gray) Crong. (Asteraceae) as their sole 
larval host plants at the study site, which are locally re- 
stricted within the treatment area. Two migratory but- 
terfly species, Danaus plexippus and Vanessa cardui 
(Scott 1986), were found in 1997 but have not be seen 
since, perhaps indicating that they too were extirpated 
from the treatment area. However, it is not known if 
there were local populations of these species in 
Schwarz Park before the spray or the individuals were 
observed migrating through the area. Considering the 
relatively small size of the spray area (32.4 ha), local 
butterfly extinction should not transpire because there 
is ample edge habitat for recolonization. However, the 
isolated nature of the spray site and the relative blan- 
keting of butterfly habitat with Bt spores suggests how 


VOLUME 56, NUMBER 3 


even common species can be at risk of local extinction 
when they are subject to pest control measures. 

In the case of Schwarz Park, the spray area was rela- 
tively small, but non-target Lepidoptera populations in 
larger spray areas may not recover as quickly as the 
Schwarz Park area. Historical Bt sprays in western 
Oregon have been large, a 120,000 ha block was re- 
peatedly sprayed in 1985, 1986, and 1987 (Johnson et 
al. 1989). With increasing evidence for the negative 
consequences of Bt on non-target Lepidoptera com- 
munities and the species that depend on Lepidoptera 
for food, the ecological impacts associated with broad- 
scale use of Bt for pest control should be questioned. 
Other alternatives that are pest species specific are the 
best option for minimizing the mortality and large- 
scale disruption of native Lepidoptera communities in 
pest eradication areas. 


I would like to thank Kat Beal for her support of this project, 
Carla M. Penz, Thomas R. Walla, and two anonymous reviewers for 
their helpful comments on this manuscript. 


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PAUL SEVERNS, U.S. Army Corps of Engineers, 
Willamette Valley Projects, PO. Box 429, Lowell, Ore- 
gon 97452, USA 


Received for publication 3 October 2001; revised and accepted 18 
April 2002. 


170 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


APPENDIX. Adult butterfly data for the Schwarz Park spray site from 1997—2000. “**” = migratory species. 


Butterfly Species Diapause Life Stage 1997 1998 1999 2000 
Hesperiidae Larvae 5 0 0 3 
Erynnis propertius (Scudder & Burgess) 
Pyrgus communis (Grote) Multivoltine 1 0 0 6 
Ochlodes s. sylvanoides (Boisduval) Eggs or Ist instar 0 0 0 
Amblyscirtes vialis (W. H. Edwards) Larvae 0 0 0 1 
Papilionidae 
Parnassius clodius claudianus Stichel Eggs 2 0 0 0 
Papilio rutulus (Lucas) Pupae 1 0) 0 9 
Papilio eurymedon (Lucas) Pupae 1 0 0 6 
Pieridae 
Pieris napi marginalis Scudder Pupae 1 0 0 3 
Pieris rapae Linnaeus Multivoltine 0 0 0 1 
Anthocharis sara flora Wright Pupae 3 1 1 3 
Colias eurytheme Boisduval Multivoltine 0 2 0 (0) 
Lycaenidae 
Mitoura grynea (Huber) Pupae 5 0 0 0 
Strymon melinus setonia McDunnough Multivoltine 8 5 5 14 
Everes comyntas (Godart) Multivoltine 5 0) 0 15 
Celastrina argiolus echo (W. H. Edwards) Pupae 13 4 4 12 
Plebejus acmon acmon (dos Passos) Multivoltine 0) 0 il 0 
Glaucopsyche lygdamus columbia (Skinner) Pupae 97 16 5D 108 
Nymphalidae 
Phyciodes mylitta mylitta (W. H. Edwards) Multivoltine 15 6 8 30 
Phyciodes p. pratensis (Behr) Pupae 1 (0) 0 0 
Boloria epithore chermocki (Perkins & Perkins) 4th instar larvae 4 (0) 0 2 
Euphydryas chalcedona colon 3rd or 4th instar larvae 2 0 0 1 
(W. H. Edwards) 
Polygonia satyrus (W. H. Edwards) Adult Il 0 2 1 
Nymphalis antiopa (Linnaeus) Adult 1 il (0) 0 
Vanessa cardui ** (Linnaeus) Adult 1 il 0 0 
Vanessa atalanta rubria ** (Fruhstorfer) Adult 1 0 0 it 
Satyridae 
Coenonympha tullia eunomia (Dornfeld) 3rd or 4th instar larvae 8 7 ll 48 
Danaidae 


bo 


Danaus plexippus ** (Linnaeus) Adult 0 0 0 


Journal of the Lepidopterists’ Society 
56(3), 2002, 171-172 


NOMENCLATURAL CORRECTNESS OF PHYCIODES PRATENSIS VS. 
PHYCIODES PULCHELLUS (NYMPHALIDAE) 


Additional key words: nomenclature, pulchella. 


Scott (1994) published an exhaustive analysis of 
Phyciodes species names, including the issue of 
whether pulchella (Boisduval, 1852) or pratensis 
(Behr, 1863) is the correct name in accordance with 
the rules of zoological nomenclature, for the wide- 
spread “Field Crescent” butterfly of western North 
America. Bird et al. (1995) adopted the use of pul- 
chella, as did Emmel et al. (1998) who emended the 
spelling to puchellus to conform to the gender congru- 
ence provisions of the International Code for Zoologi- 
cal Nomenclature (International Commission on Zoo- 
logical Nomenclature [ICZN] 1999). Other recent 
authors, such as Guppy and Shepard (2001), Layberry 
et al. (1998) and Opler (1999), used pratensis, with 
only Layberry et al. (1998) providing any reasons for 
taking this approach. This on-going disagreement re- 
garding the usage of these two names needs further 
discussion, to establish nomenclatural stability. 

Layberry et al. (1998) disagree with the use of pul- 
chella because of two alleged shortcomings in the 
original description. One stated shortcoming is that 
the name “. . . lacked a description (required for all 
new species) . . .”. This alleged shortcoming is an in- 
correct summary of nomenclatural requirements. New 
names published prior to 1931 are in fact available on 
the basis of a description, a definition or an indication 
(ICZN 1999, Article 12). An indication denotes “the 
proposal ofanew... species-group name in associa- 
tion with an illustration of the taxon being named, or 
with a bibliographic reference to such an illustration . . .” 
(ICZN 1999, Article 12.2.7). In naming pulchella, 
Boisduval (1852) provided a clear indication to two 
published illustrations that he considered to be pul- 
chella. He cited “Drury, Ins. I. PI. 21, £.5,6” which is a 
clear reference to two illustrations in Drury (1770). 
This, together with Boisduval having met the require- 
ments of ICZN (1999) Article 11, establishes the avail- 
ability of the name pulchella Boisduval, 1852. 

Scott (1994) missed the simplicity of this conclusion, 
and instead concluded that the reference to the illus- 
trations, combined with Drury’s statement that pul- 
chella is not to be confused with P. tharos or P. mor- 
pheus, constitutes a description or definition. Whether 
this conclusion is correct (it is, at best, a very poor def- 
inition) is irrelevant, because the name pulchella is 
available through an indication, regardless of whether 
a description or definition is provided. It is also inter- 
esting to note that P. tharos Drury, 1773 was named 
through indication to the same figures in Drury 


(1770). This situation gets even more interesting when 
one realises that Kirby (1837) also referenced the 
same figures in his description of P. selenis. 

The second shortcoming stated by Layberry et al. 
(1998) was that pulchella “. . . was proposed by Bois- 
duval to be applied to an illustration of Phyciodes 
tharos in Drury’s classical book (1773) . . .” and thus 
pulchella should be viewed as a junior synonym of 
tharos. While researching this issue, one colleague 
suggested that the type of pulchella is the figure in 
Drury and hence pulchella is an objective synonym of 
tharos. This cannot be. The International Code of Zo- 
ological Nomenclature (Article 72.5.6) sets out that 
the name bearing type is the specimen or specimens 
and not the illustration, although perhaps our col- 
league intended to suggest that the type was the spec- 
imen(s) on which Drury’s illustration was based. How- 
ever, Boisduval only indicated that the specimen(s) 
illustrated by Drury (1770) represented his new 
species pulchella; he did not assert that the illustration 
was the “type” specimen to which he was applying the 
name. Boisduval did not specify a holotype, and the 
actual name bearing type of pulchella is a specimen in 
the United States National Museum. The specimen 
has been designated both as a lectotype and as a neo- 
type (Scott 1994, Emmel et al. 1998). Pulchella can 
only be a subjective synonym of tharos for those 
people who might be inclined to view pulchella and 
tharos as being the same taxon. We are unaware of 
anyone who has suggested such a taxonomic interpre- 
tation. It is also irrelevant that the illustrations in 
Drury (1770) that were indicated in Boisduval’s de- 
scription of pulchella are in fact illustrations of dark 
tharos, because the type specimen determines the ap- 
plication of a name and the actual identities of all other 
specimens or illustrations are irrelevant. 

The name pulchella is therefore available in confor- 
mance with the International Code of Zoological 
Nomenclature. Furthermore, there is no question that 
the name-bearing type of pulchella is a different but- 
terfly than the name-bearing type of tharos, so pul- 
chella cannot be a synonym of tharos Drury, 1773. An 
author denying the correctness of pulchella must 
demonstrate a deficiency in the lectotype designation, 
followed by demonstration of a deficiency in the neo- 
type designation, or they stand in non-compliance with 
the International Code of Zoological Nomenclature. 
Nothing has been published to upset the correctness 
of pulchella over pratensis. So the correct name for 


this butterfly is Phyciodes pulchellus (Boisduval, 1852) 
by virtue of the provisions of the Code, including gen- 
der congruence. 


LITERATURE CITED 


Birp, C. D., G. J. HILcuie, N. G. Konpia, E. M. PIKE & F. A. H. 
SPERLING. 1995. Alberta Butterflies. The Provincial Museum 
of Alberta, Edmonton, Alberta. 349 pp. 

BorspuvaL, J. B. A. D. DE. 1852. Lepidopteres de la Californie. 
Annales Societe Entomologique France 10:275—324. 

Drury, D. 1770. Illustrations of natural history, wherein are exhib- 
ited upwards of two hundred and forty figures of exotic Insects 
_.. to which is added a translation in French. London, White 
L:xxviii + 130 + [2] pp., 50 + 1 pls, 4 f. 

EMMEL, J. F., T. C. EMMEL & S. O. Matroon. 1998. The types of 
California butterflies names by Jean Alphonse Boisduval: desig- 
nation of lectotypes and a neotype, and fixation of type locali- 
ties. Pp. 3-76 In T. C. Emmel (ed.), Systematics of Western 
North American Butterflies. Mariposa Press, Gainesville, 
Florida. 878 pp 

Gupry, C. S. & J. H. SHEPARD. 2001. Butterflies of British Colum- 
bia. UBC Press, Vancouver. 414 pp. 


Journal of the Lepidopterists’ Society 
56(3), 2002, 172-173 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE. 
1999. International Code of Zoological Nomenclature, fourth 
edition. The Natural History Museum, London, UK. 126 pp. 

Kirsy, WILLIAM. 1837. [Lepidoptera: Diurna.] Pp. 286-300, pls. 
3,4, in Part 4, Insects, In Fauna Boreali-Americana or the zool- 
ogy of the northern parts of British America, John Richardson. 
Josiah Fletcher, Norwich, England. v—xxxix + 325 pp. + cpl 1-8 
+ page i (errata). 

LayBERRY, R. A., P. W. Hatt & J. D. LAFONTAINE. 1998. The But- 
terflies of Canada. University of Toronto Press, Toronto, On- 
tario. 280. 

OpLeR, P. A. 1999. A Field Guide to Western Butterflies. 
Houghton Mifflin Co., New York, New York. 540 pp. 

Scott, J. A. 1994. Biology and systematics of Phyciodes. Papilio 
(New Series) 7:1—120. 


NorBERT G. KONDLA, Box 244, Genelle, British Co- 
lumbia VOG 1GO0 Canada, and CrisPIN S. Guppy, 4627 
Quesnel-Hydraulic Road, Quesnel, BC V2] 6P8 Canada. 


Received for publication 28 January 2002; revised and accepted 
10 May 2002. 


HEDYA SALICELLA (L.), A PALAEARCTIC SPECIES, 
COLLECTED IN NORTH AMERICA (TORTRICIDAE) 


Additional key words: immigrant, Olethreutinae, Salix, Populus. 


Fics. 1-2. Hedya salicella male from Atchison Co., Missouri. 1. Wings. 2. Genitalia (genit. slide MS 97199). 


Hedya salicella (1.) is a trans-Palaearctic species 
whose larvae feed in spun shoots and folded leaves of 
Salix and Populus species (Salicaceae). The five North 
American specimens reported here were in three dif- 
ferent collections, two public and one private. The dis- 
tinctive forewing and genitalia of these specimens 
(Figs. 1, 2) match illustrations and adult sizes in 
Bentinck & Diakonoff (1968), Bradley et al. (1979), 
and other handbooks on Eurasian Olethreutinae. They 


also match three pinned adults of H. salicella from 
England and Germany that we examined. 

The American specimens were collected over a 30- 
yr period at scattered localities: 1956 in Ontario, 1975 
in Massachusetts, and 1985 in Newfoundland and Mis- 
souri. Such a diffuse temporal-geographic pattern pro- 
vides little specific information about introduction and 
spread beyond the general conclusion that H. salicella 
is an immigrant in North America. 


VOLUME 56, NUMBER 3 


The June—August adult recovery dates are similar to 
those reported in Eurasia (Bentinck & Diakonoff 
1968, Bradley et al. 1979, Kuznetsov 1989), and sug- 
gests that H. salicella is bivoltine in North America. 
Details of wintering in Eurasia are unclear, but adult 
flight dates also suggest that the egg or partly grown 


larva is the wintering stage. 

Specimens examined. North America: Vittoria, ON, 13-VIII- 
56, Freeman & Lewis, Canadian National Collection of Arthropods 
(CNC), Ottawa, ON; Cambridge, MA, 381 Walden St. at Concord 
Ave., 26-VI-75, R. Silberglied, at 15 w blacklight, male genit. slide 
WEM 911981, forewing length 9.0 mm, Museum of Comparative 
Zoology, Harvard, University, Cambridge, MA; 2 specimens, St. 
Johns, NF, reared VII-85, CNC; Brickyard Hill Wildlife Area, Atchi- 
son Co., MO, at blacklight, J. R. Heitzman, male genit. slide MS 
97199, forewing length 9.0 mm, J. R. Heitzman collection, Inde- 
pendence, MO (Figs. 1 & 2). Europe: Derbyshire, England, 30-VI- 
25, H. C. Hayward, male genit. prep. WEM 291992, forewing 
length 9.5 mm; Mt. Kaiserstuhl, Baden-Wiirttemberg, Germany, 
6-VII-53, E. Jiickh, forewing length 10.0 mm; Capeila, Germany, 14- 
VII-42, Heddergott, male genit. prep. WEM 291991, forewing 
length 10.0 mm, all three in U. S. National Museum of Natural His- 
tory, Smithsonian Institution, Washington, DC. 


Journal of the Lepidopterists’ Society 
56(3), 2002, 173-176 


173 


We thank J. W. Brown, S. Cover, D. G. Furth, J. R. Heitzman, 
and P. D. Perkins for specimen loans. 


LITERATURE CITED 


BENTINCK, G. A., GRAAF & A. DiAKONOFF. 1968. De nederlandse 
Bladrollers (Tortricidae). Monogr. Nederl. Entomol. Ver. 3, 201 pp. 

BRADLEY, J. D., W. G. TREMEWAN & A. SmiTH. 1979. British tortricoid 
moths. Tortricidae: Olethreutinae. Ray Society, London. 336 pp. 

Kuznetsov, V. I. 1989. Leaf-rollers (Lepidoptera, Tortricidae) of 
the southem part of the Soviet Far East and their seasonal cy- 
cles, pp. 57-249. In O. L. Kryzhanovskii, (ed.), Lepidopterous 
fauna of the USSR and adjacent countries (translation). E. J. 
Brill, Leiden. 405 pp. 


MICHAEL SABOURIN. 23476 Johnson Rd., Grantsburg, 
WI 54840 USA, WILLIAM E. MILLER, Department of 
Entomology, University of Minnesota, St. Paul, Min- 
nesota 55108 USA, and P. T. Dane, Agriculture and 
Agri-food Canada, K.W. Neatby Building, Ottawa, 
Ontario, Canada KIA 0C6. 


Received for publication 14 February 1999; revised and accepted 
1 December 1999 


A HOSTPLANT EXTENSION FOR HEMILEUCA HERA HERA (SATURNIIDAE: HEMILEUCINAE): 
THREETIP SAGEBRUSH (ARTEMISIA TRIPARTITA RYDB.) 


Additional key words: sagebrush sheep moth, Artemisia tridentata, captive-rearing. 


Big sagebrush, Artemisia tridentata Nutt. (Aster- 
aceae), is widely documented as the exclusive host- 
plant for the sagebrush sheep moth, Hemileuca hera 
hera (Harris) (Ferguson 1971, McFarland 1974, 
Tuskes 1984, Stone 1991, Tuskes et al. 1996). In an in- 
vestigation of insects in the upper Snake River Plain of 
southeastern Idaho, Stafford (1987) also found larvae 
of H. hera hera only on A. tridentata hosts, despite the 
presence of A. nova Nelson, A. arbuscula Nutt., and 
A. tripartita Rydb. at the study sites. The use of 
Artemisia species other than A. tridentata has been re- 
ported only by Collins (1974), who observed a small 
number of H. hera hera larvae on silver sagebrush, A. 
cana Pursh. 

During a series of field surveys conducted to inves- 
tigate the life history of H. hera hera in the same gen- 
eral area of southeastern Idaho studied by Stafford 
(1987), I commonly found larvae on A. tridentata ssp. 
wyomingensis (Beetle & Young). However, in one 
mixed-sagebrush community, I observed larvae at all 
stages of development feeding on both A. tridentata 
ssp. wyomingensis and A. tripartita ssp. tripartita 
(hereafter abbreviated as A. tridentata and A. tripar- 
tita). In 1997 and 1998, I reared captive larvae on A. 
tripartita for the purpose of obtaining voucher speci- 


mens. My observations of host associations and the re- 
sults of captive-rearing are discussed in this note. 

Field-observations. During April 1997, an inten- 
sive search was conducted of the area in which H. hera 
hera larvae were observed feeding on A. tridentata 
and A. tripartita hosts. Plants containing larvae were 
tagged and larval development was monitored weekly 
between May 1997 and July 1997. Larvae on both A. 
tridentata and A. tripartita demonstrated a life history 
and behavior typical of H. hera hera in other parts of 
its range. On both hosts, first through third instars gen- 
erally fed gregariously and then dispersed to feed indi- 
vidually as fourth and fifth instars. On multiple occa- 
sions, late-instar larvae were observed to move from 
one Artemisia species to the other. Larvae often re- 
mained on the second host for several days, indicating 
that plants of both species were acceptable food 
sources. 

Female H. hera hera in this study area also used 
both A. tridentata and A. tripartita as oviposition hosts 
(Hampton 2000). Eggs were most commonly laid on 
stems of both Artemisia species, but approximately 
18% were located on the stems of other species in the 
sagebrush understory including Chrysothamnus vis- 
cidiflorus (Hook.) Nutt. (Asteraceae), Leptodactylon 


174 


pungens (Torr.) Nutt. (Polemoniaceae), and Bromus 
tectorum L. (Poaceae) (Hampton 2000). Despite these 
apparent oviposition “mistakes,” feeding by larvae on 
plants other than Artemisia spp. was observed only 
once. In the spring of 1997, several larvae developed 
to the third instar while feeding on green rabbitbrush 
(C. viscidiflorus). However, compared to larvae feed- 
ing on nearby sagebrush, these larvae were severely 
stunted. Continued searches of this plant showed no 
evidence that the larvae matured or pupated. 

Previously documented instances of H. hera hera 
feeding on plants of genera other than Artemisia are 
also rare. Collins (1974) reported a single H. hera hera 
larva feeding on lupine (Lupinus sp., Fabaceae) and 
use of buckwheat (Eriogonum spp., Polygonaceae) has 
been reported (Tuskes 1984). These observations have 
been generally attributed to “adventitious” feeding or 
conditions associated with larval overcrowding (Collins 
1974, Tuskes 1984). 

Captive-rearing. During late June and early July 
of 1997, I collected 31 fourth and fifth instar H. hera 
hera larvae from plants of both A. tridentata and A. 
tripartita for the purpose of obtaining adult voucher 
specimens (archived in the entomology collection at 
Idaho State University, International Collection Reg- 
istry ICIS). 

Larvae were fed on potted A. tripartita plants lo- 
cated in screened outdoor cages. The cages were con- 
structed of rigid wire mesh rolled into free-standing 
cylinders approximately 0.60 m in diameter. The cylin- 
ders were covered with 1 mm mesh nylon screen, 
placed over potted sagebrush plants resting on soil- 
filled bases of equal diameter, and pressed into the soil 
to form an escape-proof seal. By 21 July 1997, 22 of 
the 31 caterpillars had successfully pupated. On 10 
November 1997, the soil-covered pupae were moved 
to an unheated garage where they were left in dia- 
pause. In April 1998, the cages were reconstructed 
and the pupae were returned to the outdoors to await 
eclosion. Five adult H. hera hera (3 females and 2 
males) emerged between 11 and 26 August 1998 (see 
Table 1). In early September 1998, the soil bases of all 
cages were excavated and four potentially viable pupae 
were transferred to small containers and moved in- 
doors. Two of the pupae in containers subsequently 
produced adults, one female on 12 September 1998, 
and a male on 2 October 1998. Of the 22 pupae pro- 
duced in 1997, 15 were lost to various causes including 
parasitism by a fungus and an unidentified wasp (Ich- 
neumonidae: Hymenoptera) (Table 1). 

In April 1998, a group of first instar larvae collected 
from A. tridentata were reared in the same manner, 
using the same cages and A. tripartita plants. Cold, 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


TABLE 1. Summary of captive-rearing results. 


1997-1998 1998-1999 
No. of initial larvae 318 95» 
No. of pupae produced 22 (71%) T° (28%) 
No. of adults produced? 7* (23%) 3° (12%) 
No. of unsuccessful pupae 15° (48%) 4° (16%) 
Damaged in handling a = 
Fungal parasites 7 3 
Ichneumonid parasitoid 3 — 
Other insect damage 1 = 
Unknown causes 3 1 


“Fourth and fifth instars collected between June and July 1997. 

> Estimated number of first instars collected as an aggregation in 
April 1998. 

© All but seven larvae were killed by cold, wet weather conditions 
that occurred as larvae were molting from second to third instars. 

‘Dates of emergence for seven 1998 adults: d 11 Aug., 2 23 August, 
d 24 Aug., 2 26 Aug., 2 27 Aug., 2 12 Sept.!, and 6 2 Oct! Dates of 
emergence for three 1999 adults: 9 3 Aug., d 8 Aug., and 2 9 Aug. 

©Total based on the assumption that adults emerged from 1997 pu- 
pae (see item “f” below). 

‘Late dates of adult eclosion may indicate same-year emergence. 


wet weather killed all but seven of the approximately 
25 larvae during their second molt. However, the size 
and vigor of the remaining larvae appeared similar to 
larvae observed in the field and all seven pupated by 
late July 1998. When cages were dismantled in early 
September 1998, three pupae had been destroyed by a 
fungal parasite. Four undamaged pupae were re- 
moved from the soil to small containers and refriger- 
ated during November 1998 through March 1999. 
Three adults (1 male and 2 females) emerged between 
3 and 9 August 1999 (Table 1). However, two of these 
individuals were dark-colored, appeared weak, and 
the wings of both never fully expanded. 

The potential for alternative hosts. Fourteen 
species of Artemisia and 11 subspecies in the subgenus 
Tridentatae are found within the western United States 
(McArthur et al. 1998, McArthur 2000). Although A. 
tridentata and its subspecies have the greatest geo- 
graphic range (McArthur 2000) and are found through- 
out the range of H. hera hera (Fig. 1), the distributions 
of most members of the subgenus intersect the moth’s 
range to some extent. The northeastern range of H. 
hera hera extends well outside the range of A. tridentata 
(Fig. 1) and beyond the ranges of all Artemisia species 
except A. cana (McArthur & Plummer 1978), indicating 
the potential for an alternate host in that area. 

Artemisia tripartita ssp. tripartita is generally re- 
stricted to areas of Idaho, Washington, and Oregon 
(Shultz 1984), and occurs only in the central portion of 
the range of H. hera hera (Fig. 1). A second subspecies, 
A. tripartita ssp. rupicola Beetle, inhabits large areas of 
Wyoming (Beetle 1960, Shultz 1984). Although plant 
morphology differs markedly between A. tripartita and 


VOLUME 56, NUMBER 3 


A. tridentata, early leaf growth in both species (Miller 
& Shultz 1987) coincides with larval emergence (pers. 
obs.), and chemical constituents of both are similar 
(Kelsey & Shafizadeh 1979). However, differences in 
plant structure are known to influence microclimatic 
conditions within and around plants (Hinds & Rickard 
1968, Lawton 1983, Strong et al. 1984, Pierson & 
Wight 1991) and can impact selection of oviposition 
site, larval survivorship and development, and preda- 
tion dynamics in H. hera hera (Hampton 2000). Unlike 
most woody Artemisia species, A. tripartita plants can 
re-sprout from the roots after fire, over-grazing, or win- 
ter damage (Beetle 1960, Hironaka et al. 1983). Be- 
cause A. tridentata plants are easily killed by fire (Bee- 
tle 1960) and take many years to regenerate from seed 
(Hironaka et al. 1983), the use of A. tripartita as an al- 
ternative host could have important implications with 
regard to long-term survival and viability of H. hera 
hera populations. This may be especially important in 
the wake of more frequent wildfire and other large- 
scale disturbances to sagebrush communities across the 
Great Basin (Knick & Rotenberry 1997, Knick 1999). 

Another subspecies of H. hera is also known to use 

more than one hostplant. Both A. tridentata and A. fili- 

folia Torr. are used as hosts by H. hera magnifica (Rot- 
ger), depending on the location within its range (Stone 
& Smith 1990, Stone et al. 1988). Because Artemisia 
species and subspecies share many chemical similarities 
(Kelsey & Shafizadeh 1979), it may be reasonable to ex- 
pect that other species might also serve as suitable food- 
plants for H. hera hera. Nevertheless, some herbivorous 
insects demonstrate apparent preferences for certain 
subspecies and hybrids of A. tridentata (Messina et al. 
1996) and there is evidence that some Artemisia species 
and varieties can be unsuitable hosts for H. hera hera 
larvae (McFarland 1974, Tuskes et al. 1996). The varia- 
tion in secondary chemistries of Artemisia species and 
hybrids (McArthur et al. 1988, Messina et al. 1996) may 
influence the selection of hosts by H. hera hera. Al- 
though hybridization between A. tridentata ssp. 
wyomingensis and A. tripartita is a possibility, it has not 
been documented (B. L. Welch, pers. com.) and at- 
tempts to produce hybrids under controlled conditions 
have been unsuccessful (McArthur et al. 1998). 

The use of other species and varieties within the 
genus Artemisia by H. hera hera may simply not appear 
more frequently in the literature because sagebrush 
identification and taxonomy is complicated. Prior to 
1960, several species were included as subspecies within 
the A. tridentata complex (Hall & Clements 1923), but 
were assigned species status in later treatments (Beetle 
1960, McArthur et al. 1998). For example, A. tripartita 
was formerly classified as A. tridentata ssp. trifida Hall & 


175 


@ 7 herahera (Tuskes et al. 1996) 
=== Artemisia tridentata (McArthur 2000) 
#*= Artemisia tripartita (McArthur and Plummer 1978) 


Fic. 1. The ranges of Hemileuca hera hera and Artemisia host- 
plants. 


Clements (Hall & Clements 1923). Although leaf and 
plant morphologies of A. tripartita are distinctive, host- 
plant nomenclature to the level of subspecies is rarely, if 
ever, included in published food records for H. hera 
hera. Consequently, some accounts could refer to a 
number of Artemisia species formerly designated as sub- 
species of A. tridentata. 

The use of A. tripartita by H. hera hera in southeast- 
em Idaho provides new evidence that the sagebrush 
sheep moth can successfully exploit hosts other than A. 
tridentata in some parts of its range. The potential use of 
alternative Artemisia hosts by H. hera hera in other geo- 
graphical locations, including A. cana in the northeastern 
extension of its range, warrants further investigation. 


I am grateful to Paul Severns, Stefan Sommer, and Paul Castro- 
villo for comments and suggestions that improved this manuscript. I 
also thank Jacalyn Brower for her assistance in developing the host 
range map. 


LITERATURE CITED 


BEETLE, A. A. 1960. A study of sagebrush: The section Tridentatae 
of Artemisia. Bulletin No. 363, Wyoming Agricultural Experi- 
ment Station, University of Wyoming, 83 pp. 

Couns, M. M. 1974. The population dynamics of moths in the 
genus Hemileuca on Monitor Pass, California. Unpublished 
master’s thesis, California State University, Hayward, California. 

FERGUSON, D. C. 1971. Bombycoidea: Saturniidae. Moths of North 
America. Fascicle 20.2A. E. W. Classey Ltd., London. 153 pp. 

HALL, H. M. & F. E. CLEMENTS. 1923. The phylogenetic method 
in taxonomy: the North American species of Artemisia, 
Chrysothamnus, and Atriplex. Publ. Carnegie Inst. Wash. 
326:1—355. 

Hampton, N. L. 2000. Hostplant selection and egg placement as 
factors influencing egg survival in Hemileuca hera hera (Sat- 
urniidae: Lepidoptera). Unpublished master’s thesis, Idaho 
State University, Pocatello, Idaho. 

Hinps, W. T. & W. H. Rickarp. 1968. Soil temperatures near a 
desert steppe shrub. Northwest Sci. 42(1):5-13. 


176 


Hrronaka, M., M. A. FosBerc & A. H. WINWARD. 1983. Sagebrush- 
grass habitat types of southern Idaho. University of Idaho Forest, 
Wildlife, and Range Bulletin, No. 35, Moscow, Idaho. 44 pp. 

KELSEY, R. G & F. SHAFIZADEH. 1979. Sesquiterpene lactones and 
systematics of the genus Artemisia. Phytochem. 18:1591-1611. 

KNICK, S. T. 1999. Requiem for a sagebrush ecosystem? Northwest 
Sci. 73(1):53-57. 

Knick, S. T. & J. T. ROTENBERRY. 1997. Landscape characteristics 
of disturbed shrubsteppe habitats in southwestern Idaho 
(U.S.A.). Landscape Ecol. 12:287-297. 

Lawton, J. H. 1983. Plant architecture and the diversity of phy- 
tophagous insects. Ann. Rev. Entom. 28:23-39. 

McArtuur, E. D. 2000. Sagebrush systematics and distribution. 
Pages 9-14. In P. G. Entwistle, A. M. Debolt, J. H. Kaltenecker 
& kK. Steenhof (compilers). Proceedings: Sagebrush Steppe 
Ecosystems Symposium. Bureau of Land Management Publi- 
cation No. BLM/ID/PT-001001 + 1150, Boise, Idaho, U.S.A. 

McArtHur, E. D. & A. P. PLUMMER. 1978. Biogeography and 
management of native western shrubs: A case study, section Tri- 
dentatae of Artemisia. Intermountain Biogeography: A Sympo- 
sium. Great Basin Nat. Memoirs. 2:229-243. 

McARTHUR, E. D., B. L. WELCH & S. C. SANDERSON. 1988. Nat- 
ural and artificial hybridization between big sagebrush 
(Artemisia tridentata) subspecies. J. Heredity 79:268-276. 

MCARTHUR, E. D., J. MUDGE, R. VAN BUREN, W. R ANDERSON, S. C. 
SANDERSON & D. G. BABBEL. 1998. Randomly amplified poly- 
morphic DNA analysis (RAPD) of Artemisia subgenus Triden- 
tatae species and hybrids. Great Basin Nat. 58:12-27. 

McFarLANp, N. 1974. Notes on three species of Hemileuca (Sat- 
urniidae) from eastern Oregon and California. J. Lepid. Soc. 
28:136-141. 

MESSINA, F- J., J. H. RICHARDS & E. D. McARTHUR. 1996. Variable 
responses of insects to hybrid versus parental sagebrush in a 
common garden. Oecologia 107:513-521. 

MILLER R. F. & L. M. Suurz. 1987. Development and longevity 
of ephemeral and perennial leaves on Artemisia tridentata 
Nutt. ssp. wyomingensis. Great Basin Nat. 47(2):227—230. 


Journal of the Lepidopterists’ Society 
56(3), 2002, 176-178 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Pierson, F. B. & J. R. WicuT. 1991. Variability of near-surface 
temperature on sagebrush rangeland. J. Range Man. 
44:49]—497. 

SHULTZ, L. M. 1984. Taxonomic and geographic limits of Artemisia 
subgenus Tridentatae (Beetle) McArthur (Asteraceae: An- 
themideae). Pages 20-28. In E. D. McArthur, B. L. Welch 
(compilers). Proceedings—Symposium on the Biology of 
Artemisia and Chrysothamnus, July 9-13 1984, Provo Utah. 
Genera Technical Report INT-200, USDA, Forest Service, In- 
termountain Research Station, 1986, 398 pp: 

STAFFORD, M. P. 1987. Insect interactions with four species of 
sagebrush (Artemisia) in southeastern Idaho. Unpublished doc- 
toral dissertation, University of Idaho, Moscow, Idaho. 

STONE, S. E. 1991. Foodplants of world Saturniidae. Memoir 
Number 4, The Lepidopterists’ Society. 

STONE, S. E. & M. J. SmitH. 1990. Buckmoths (Lepidoptera: Sat- 
urniidae: Hemileuca) in relation to southwestern vegetation and 
foodplants. Desert Plants 10 (1):13-30. 

STONE, S. E., D. E. Swirt & R. S. PEIGLER. 1988. The life history 
of Hemileuca magnifica (Saturniidae) with notes on Hemileuca 
hera marcata. dl Res. Lepid. 26(1-4):225-235. 

STRONG, D. R., J. H. Lawron & R. SouTHWooD. 1984. Insects on 
plants, community patterns and mechanisms. Harvard Univer- 
sity Press, Cambridge, Massachusetts. 

TuskeEs, P. M. 1984. The biology and distribution of California 
Hemileucinae (Saturniidae). J. Lepid. Soc. 38:281-309. 

Tuskes, P. M., J. P. TurrLe & M. M. Coins. 1996. The wild silk- 
moths of North America: A natural history of the Satuniidae of 
the United States and Canada. Cornell Univ. Press, Ithaca, New 


York. 250 pp. 


NANcy HAMPTON, 545 E. 13th Street, Idaho Falls, 
Idaho, 83404, USA 


Received for publication 10 September 2001; revised and ac- 
cepted 5 March 2002. 


REDISCOVERY OF AND NOTES ON XANTHOTHRIX RANUNCULI 
FORM ALBIPUNCTA BARNES & BENJAMIN, 1925 (NOCTUIDAE: STIRIINAE) 


Additional key words: discal spot, Coreopsis. 


Barnes and Benjamin (1925) described albipuncta 
as a form of Xanthothrix ranunculi based on two indi- 
viduals, a male holotype and a female allotype, col- 
lected 10 April 1902 at Kaweah, Tulare Co., California. 
The moths were described as having “head, thorax and 
forewing black dusted with olivaceous, the latter with 
a round white spot near end of cell. Secondaries black- 
ish.” 

Xanthothrix ranunculi was described by Henry Ed- 
wards (1878) from Havilah, Kern Co., California, and 
most specimens seen are from Lovejoy Buttes, Los 
Angeles Co., California. These moths were described 
is, “Primaries wholly rich buff, sometimes pale orange 
when very fresh, but upon some scales being removed, 
showing a blackish tint. Secondaries, dusky along the 
costa, apical margin and at base, buff in the center, and 
towards the anal angle. . .” Comstock and Henne 


(1940) described their early stages, and reported the 
hostplant is Coreopsis douglasii (Asteraceae). Poole 
(1994) makes no mention of form albipuncta. 

On 21 March 1990, I collected three individuals of a 
small moth flying near Coreopsis stillmanii (Asteraceae) 
on a steep, south-facing slope above the middle fork of 
the American River, 13 mi. NE of Auburn, Placer Co., 
California. One, a worn specimen, fit the description of 
Xanthothrix ranunculi form albipuncta; the two fresh 
specimens had gold forewings with a light yellow spot 
near the end of the cell. The habitat consisted of annual 
grasses and flowering plants surrounded by foothill 
woodland. At the same locality, in March 1998 the 
moths were numerous and were often seen sitting on 
the Coreopsis flower heads. More individuals were col- 
lected and the association with Coreopsis observed. 
Alvin Ludtke collected females and Coreopsis still- 


VOLUME 56, NUMBER 3 


Fic. 1. Xanthothrix ranunculi form albipuncta. Columns 1 and 2, Mariposa Co., showing variation. Column 1| top, Scotch Gulch, 6 mi. SE 
of Coulterville, Mariposa Co., CA, 29-III-1999, others, same locality but 22-11-2000. Column 3 Placer Co. 2 mi. E of Ruck-A-Chucky Falls, 
middle fork of the American River, 13 mi. NE of Auburn, Placer Co., CA. Top 27-III-1999, bottom 19-III-1998. 


Fic. 2. Mature larva of Xanthothrix ranunculi form albipuncta 
on Coreopsis stillmanii. Reared ex ovum from a female collected 2 
mi. E of Ruck-A-Chucky Falls, middle fork of the American River, 
13 mi. NE of Auburn, Placer Co., CA. Scale at top of photograph in 
millimeters. 


manni from the site to attempt rearing. The females 
oviposited on these plants, and larvae (which fed on the 
developing achenes) were reared through the third in- 
star. The larvae were then moved to a perennial g garden 
Coreopsis, on which they fed for a while, then died, for 
unknown reasons, without pupating. 

In March 1999 and March 2000, D. Brown and I 
collected these moths southeast of Coulterville, Mari- 
posa Co., California resting on the flower heads of 
Coreopsis stillmanii. Goldfields, Lasthenia (Aster- 
aceae) were growing with the Coreopsis there but the 
moths did not rest on their flower heads. Unlike in 
Placer Co., the habitat there was on serpentine soils, 
and Buck Brush, Ceanothus cuneatus, dominated the 
flora. These moths very closely resembled the Placer 
Co. moths, although a greater number of them had a 
dull, gray-green color to the forewing, with the 
forewing spot closer to white. This coloring was most 
marked in worn individuals, but was also characteristic 
of some fresh individuals, on which the long scales on 
the forewing were a very light yellow rather than gold. 
No individuals were found in an April 1999 visit to the 
type locality. On March 18, 2001 the moths were 
found, again associated with Coreopsis stillmanii, on 
an area of serpentine soils in Tuolumne Co. 


The Placer and Mariposa Co. moths appear more 
golden than the worn type of albipuncta, suggesting 
that the gold scaling on the forewings is gradually lost 
causing the wings to appear olivaceous in older speci- 
mens. All Mojave Desert specimens of ranunculi lack 
the yellowish spot near the apex of the discal cell that 
is present in albipuncta. 

It is likely that albipuncta and ranunculi are differ- 
ent species. Xanthothrix ranunculi is golden yellow, is 
apparently restricted to the Mojave Desert, uses Core- 
opsis douglasii as a food plant, and lacks a forewing 
spot near the apex of the discal cell. Xanthothrix ra- 
nunculi form albipuncta has golden forewings and 
blackish hindwings, is apparently restricted to the 
lower, grassy foothills of the Sierra Nevada, uses Core- 
opsis stillmanii as a food plant, and has a light yellow 
forewing spot near the apex of the discal cell. 
Philotiella speciosa and its subspecies bohartorum (Ly- 
caenidae), which also may be a separate species, dis- 
play a similar disjunct distribution. 

Plants were identified using Hickman (1993). 


I thank Alvin Ludtke, North Highlands, California, for assis- 
tance in the field, for rearing efforts, for the photograph of the 
larva, and for many discussions about these moths; Tim McCabe, 
New York State Museum, for advice on the identity of the moths; 


Journal of the Lepidopterists’ Society 
56(3), 2002, 178-179 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Steve Heydon, University of California, Davis, for obtaining the 
Henry Edwards paper, and for taking the photograph of the adults; 
and John DeBenedicits, University of California, Davis for sugges- 
tions which greatly improved this note. I also thank my wife, Doris 
Brown, for companionship and help in the field. 


LITERATURE CITED 


Barnes, W. & F. H. BENJAMIN. 1925. New and rare Lepidoptera 
from the Southwest. Pan-Pac. Entolom. II:12—15. 

Comstock, J. A. & C. HENNE. 1940. Notes on early stages of Xan- 
thothrix ranunculi. Bull. So. Calif. Acad. Sci. 9:198—199. 

Epwarps, H. 1878. Pacific Coast Lepidoptera, No. 29. Description 
of some new genera and species of Noctudiae. Proc. Calif. 
Acad. of Sci. 29:7. 

HICKMAN, J. (ed.). 1993. The Jepsen Manual. Higher Plants of Cal- 
ifornia. University of California Press. 1400 pp. 

PooLe, R. W. 1994. Noctuoidea, Noctuidae (Part). In R. B. Do- 
minick et al., The Moths of North America north of Mexico, 
fasc. 26.1:113-114. 

Prigstar, R. C. & J. F. EMMEL. 1998. An extraordinary new sub- 
species of Philotiella speciosa (LEPIDOPTERA: LY- 
CAENIDAE) from Coastal Santa Barbara County, California. 
Pg. 283-284 In T. C. Emmel (ed.), Systematics of Western 
North American Butterflies. Mariposa Press. 878 pp. 


WILLIAM D. PATTERSON, 2624 4th Avenue Sacra- 
mento, California 95818, USA 


Received for publication 19 January 2001; revised and accepted 3 
December 2001. 


ADDENDUM TO THE SPHINGIDAE OF LOUISIANA 


Additional key words: _ bait traps, hawkmoths, light traps, Louisiana, sphinx moths. 


The Sphingidae of Louisiana were reported by Brou 
and Brou (1997) in a 26-year study treating abun- 
dance, distribution, and flight periods. We listed 55 
species of Sphingidae recorded for Louisiana and 46 
species documented by our study. The quantity of 
adult sphingidae captured from 1970 to 1995 for the 
state of Louisiana totaled 71,836 specimens. This brief 
article is intended to add to and finalize our previous 
investigations. No newly recorded species were en- 
countered during these four years, nor were species 
newly encountered in bait traps versus light traps than 
were previously reported. This addendum, represent- 
ing four years (1996-1999) yielded 12,053 specimens 


representing 36 species (Table 1). Fermenting bait 
traps were operated only in 1996-1997, while ultravio- 
let light traps were operated all four years. These addi- 
tional records represent 30 consecutive years of Sphin- 
gidae records for the state of Louisiana, totaling 83,889 
captured adults. 

Total trap hours expended during the 30 year inves- 
tigation were in excess of 1.4 million hours, involving 
491,000 ultraviolet light trap hours and 913,000 fer- 
menting bait trap hours. Specimens retained during 
this addendum period are deposited in Florida State 
Collection of Arthropods (Gainesville) and Louisiana 
State University (Baton Rouge). 


VOLUME 56, NUMBER 3 


TABLE 1. 


tween 1996 and 1999. 


co 


ID UB wp 


Species 


. Agrius cingulata (F.) 


Manduca sexta (L.) 


. M. quinquemaculata (Haw.) 

. M. rustica (F.) 

. M. jasminearum (Guer.) 

. Dolba hyloeus (Drury) 

. Ceratomia amyntor (Geyer) 

. C. undulosa (Wlk.) 

. C. catalpae (Bdv.) 

. C. hageni (Grt.) 

. Isoparce cupressi (Bdv.) 

. Paratrea plebeja (F.) 

. Sphinx eremitus (Hbn.) 

. S. leucophaeta Clem. 

. S. chersis (Hbn.) 

. S. franckii Neum. 

. S. kalmiae J. E.Smith 

. S. drupiferarum J. E.Smith 

. Lapara coniferarum (J. E.Smith) 
. L. phaeobrachycerous Brou 

. Smerinthus jamaicensis (Drury) 
. Paonias excaecatus (J. E.Smith) 
. P. myops (J. E.Smith) 

. P. astylus (Drury) 

. Laothoe juglandis (J. E.Smith) 

. Pachysphinx modesta (Harr.) 
DT, 
. Erynnis alope (Drury) 
. E. lassauxi (Bdv.) 

. E. ello (L.) 

. E. obscura (F.) 

32. 
. Pachylia ficus (L.) 

. Aellopos titan (Cram.) 

. A. fadus (Cram.) 

. Enyo lugubris (L.) 

. Hemaris thysbe (F.) 

. H. diffinis (Bdv.) 

. Eumorpha satellita licaon (Cram.) 
. E. pandorus (Hbn.) 

. E. intermedia (B. P.Clark) 

. E. achemon (Drury) 

. E. vitis (L.) 

. E. fasciatus (Sulz.) 

. E. lubruscae (L.) 

. Sphecodina abbottii (Swainson) 

. Deidamia inscripta (Harr. ) 

. Amphion floridensis B. P.Clark 

. Proserpinus gaurae (J. E.Smith) 
. Darapsa versicolor (Harr.) 

. D. myron (Cram.) 

. D. pholus (Cram.) 

. Xylophanes pluto (F.) 

. X. tersa (L.) 

. Hyles lineata (F-.) 


Pseudosphinx tetrio (L.) 


E. domingonis (Btl.) 


Total records 


List of species and number of individuals collected be- 


Number of adults 


95 
36 


bo 
Soe 
He CO 


ee 


po 
bo iw) co ~l 
SOMAiooeoranoooeeEWWYH © SC Se SO KX @ 


179 


LITERATURE CITED 


BrOU, VERNON A. & C. D. Brou. 1997. Distribution and phenolo- 
gies of Louisiana Sphingidae. J. Lepid. Soc. 51:156-175. 


VERNON A. BROU JR. AND CHARLOTTE D. BrROU, 
74320 Jack Loyd Road, Abita Springs, Louisiana 
70420 USA Email: vabrou@bellsouth.net 


Received for publication 15 October 2001; revised and accepted 
26 February 2002. . 


Journal of the Lepidopterists’ Society 
56(3), 2002, 180-181 


THE CORRECT SPELLINGS AND TYPE LOCALITIES OF 
BUTTERFLIES NAMED FROM MOUNT RAINIER, WASHINGTON, USA 


Additional key words: Boloria, Clossiana, Speyeria, chariclea, titania, rainieri, mormonia, bischoffii. 


The original description of the taxon Brenthis char- 
iclea ranieri Barnes and McDunnough, 1913 is titled 
“B. CHARICLEA RAINIERI subsp. nov. (PI. II. Figs. 1—4).” 
The figure captions read: “Fig. 1. Brenthis chariclea 
ranieri B. & McD. Paradise Valley, Mt. Ranier. J Co- 
type.”; “Fig. 2. Brenthis chariclea ranieri B. & McD. 
Paradise Valley, Mt. Ranier. ° Type.”; “Fig. 3. Brenthis 
chariclea ranieri B. & McD. Paradise Valley, Mt. 
Ranier. d underside.”; and “Fig. 4. Brenthis chariclea 
ranieri B. & McD. Paradise Valley, Mt. Ranier. ° un- 
derside.” (Barnes & McDunnough 1913). The sub- 
species name is not otherwise mentioned in the origi- 


nal description. Therefore there are two spellings of 


the new subspecies in the original description, rainieri 
and ranieri, and the correct spelling must be deter- 
mined through application of the International Code 
of Zoological Nomenclature (International Commis- 
sion on Zoological Nomenclature 1999). 

Statements in the original description include 
“Hasirar. Mt. Ranier, Wash. (July 24-31) (6-7000 ft.) 
(McDunnough), 7 d, 7 2. Types. Coll. Barnes.” and 
“The species was the commonest butterfly collected, 
being found all over the grassy slopes around the so- 
called Paradise Valley.” Immediately prior to the origi- 
nal description, on the same page, Barnes and Mc- 
Dunnough state “Two years ago we captured a long 
series of specimens on Mt. Ranier, Washington. . . .”, 
and on the previous page of the publication “Mt. 
Ranier” is given as the type locality of Argynnis 
bischofft washingtonia Bames and McDunnough. It is 
therefore clear that the type locality is Paradise Valley, 
Mt. Ranier, Washington. It is also unambiguously clear 
that the subspecies was named after Mt. Ranier. 

The International Code of Zoological Nomenclature 
(International Commission on Zoological Nomencla- 
ture 1999) states: 


“32.5. Spellings that must be corrected (incorrect original 
spellings). 

32.5.1. If there is in the original publication itself, without re- 
course to any external source of information, clear evidence of an in- 
advertent error, such as a lapsus calami or a copyist’s or printer's er- 
ror, it must be corrected.” 

and 

“Examples. If an author in proposing a new species-group were 
to state that he or she was naming the species after Linnaeus, yet the 
name was published as ninnaei, it would be an incorrect original 
spelling to be corrected to linnaei.” 

and 

“33.2.2. The correction of an incorrect original spelling in accor- 
dance with Article 32.5 is a “justified emendation,” and the name 
thus corrected retains the authorship and date of the original 
spelling [Art. 19.2]. 


33.2.3. Any other emendation is an “unjustified emendation’”. . . . 
[Article 33.2.3 continues with conditions under which “prevailing 
usage” requires use of an emendation even if it was unjustified. | 


Prevailing usage (Art. 33.2.3) is clearly only a factor 
if article 32.5 does not apply. Articles 32.2.1 and 32.2.2 
refer to the choice of correct spelling by a first reviser 
and are only pertinent when the incorrect spelling is 
not demonstrably incorrect through article 32.5, which 
is not the case in this example. Since article 32.5 ap- 
plies, because it is unambiguous that the subspecies 
was named after Mt. Ranier, the incorrect spelling 
rainieri in the title of the original description must be 
corrected to ranieri. Miller and Brown (1981, taxon 
589b and note 475) were incorrect in emending the 
spelling of the name to rainieri. 

We have also reviewed topographical maps and 
other information to confirm the correspondence of 
the stated type locality data to modern information. 
Paradise Valley is a flat-bottomed mountain valley on 
the south side of Mt. Rainier through which runs Par- 
adise River. This is now the most visited location in the 
park, with abundant alpine flowers in the summer and 
deep (6 meter) snow in the winter (Reese 2001). The 
flat bottom of Paradise Valley is at 5000-5100 feet ele- 
vation. The original description states that ranieri was 
“found all over the grassy slopes around the so-called 
Paradise Valley,” indicating that the true elevation at 
which the type series was collected may have been 
56000 feet. The steep slopes at the upper (north) end 
of the valley, from which the headwaters of the Par- 
adise River originate, rise above 6000 feet to Mt. 
Rainier. Hence, for lack of any strong evidence to the 
contrary, we recommend leaving the stated elevation 
of 6-7000 feet unchanged. The boundary between 
Pierce County and Lewis County almost exactly bi- 
sects Paradise Valley. The type specimen of ranieri 
may have originated from either County. The type lo- 
cality of Brenthis chariclea ranieri Barnes and Mc- 
Dunnough, 1913 should be therefore be cited as “Par- 
adise Valley, Mt. RaliJnier, [elevation] 6—7000 ft., 
[Pierce or Lewis County], Wash[ington, USA],” with 
the square brackets indicating extrapolated data. 

Barnes and McDunnough (1913) also described the 
subspecies Argynnis bischoffi washingtonia (now re- 
ferred to as Speyeria mormonia washingtonia), with a 
similar type locality to ranieri. The differences in type lo- 
cality citation were in the elevation (7000 ft.) and refer- 
ence to only Mt. Ranier and not Paradise Valley. This in- 
dicates that the type series of washingtonia was probably 
collected on the steep slopes above the upper (north) 


VOLUME 56, NUMBER 3 


end of Paradise Valley, within Pierce County. The type 
locality of Argynnis bischofft washingtonia Barnes and 
McDunnough, 1913 should therefore be cited as “Mt. 
RaliJnier, [elevation] 7000 ft., [Pierce County], Wash|in- 
gton, USA],” to reflect accurately the information in the 
original description. It is worth noting that the spelling 
bischoffvi is the original spelling (Edwards 1870) and 
Barnes and McDunnough’s (1913) spelling of bischoffi 
was incorrect (International Commission on Zoological 
Nomenclature 1999, Article 31.1). 

Gunder (1932) named “Argynnis eurynome [=Sper = 
eria mormonia] washingtonia B. & McD., form 
ranierensis” from specimens collected at “Paradise 
Valley, Mount Ranier, Pierce Co., Washington, Aug. 
12, 1931.” Miller and Brown (1981) incorrectly and 
without explanation emended the name of this form to 
rainierensis. The correct spelling is form ranierensis, 
because it is unambiguously clear that the form was 
named after Mount Ranier. The type locality for form 
ranierensis should be cited as “Paradise Valley, Mount 
RaliJnier, Pierce Co[unty], Washington, [USA].” 

Mount Rainier, elevation 14,411 feet, is in the center 
of Mount Rainier National Park. Captain George Van- 
couver named it in 1792 to honor Rear-Admiral Peter 
Rainier of the Royal Navy (Reese 2001). On 2 March 
1899 President McKinley signed into law the bill that 
created a new national park around Mount Rainier. The 
national park was named after the mountain, but was 
misspelled “Mount Ranier National Park” in the bill. An 
image of the original bill, showing the misspelling, can 
be viewed on the Library of Congress website at 
http://Ieweb2.loc.gov/law/GLINv1/GLIN. html. The of- 
ficial, but misspelled, name may well have been used on 


181 


signs in the park in 1911, when the specimens of ranieri 
were collected, and in 1931 when the types of ranieren- 
sis were collected. We made contacted the National 
Parks Service, but could not determine when, if ever, 
the name of the national park was changed to “Mount 
Rainier National Park”. However, it appears that up un- 
til at least 1931 the spelling “Mount Ranier” was a valid 
and widely used alternate spelling for Mount Rainier. 
This is the origin of the spelling Mount Ranier as the 
type locality for the two butterfly names ranieri Bames 
and McDunnough, 1913 and ranierensis Gunder, 1932. 


LITERATURE CITED 


BARNES, WILLIAM & J. H. MCDUNNoUGH. 1913. New N. Am. Lep- 
idoptera with notes on described species. Contr. nat. hist. Lep. 
N. Amer. 2(3):91-164, pls 1-9. Decatur, Illinois: Review Press. 

Epwarps, W. H. 1870. Descriptions of new North American diur- 
nal Lepidoptera. Trans. Ent. Soc. Lond. 3(2):189-196. 

GUNDER, J. D. 1932. New Rhopalocera (Lepidoptera). Can. Ent. 
64(12):276-284. 

INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE. 
1999. Intemational code of zoological nomenclature, fourth 
edition. London: The Intemational Trust for Zoological 
Nomenclature. xxix + 306 pp. 

Miter, L. D. & F. M. Brown. 1981. A catalogue/checklist of the 
butterflies of America north of Mexico. Lepid. Soc. Mem. 
2:i-vii, 1-280. 

REESE, G. F. 2001. Washington Place Names. Tacoma, Washing- 
ton: Tacoma Public Library website, 
http://Awww.tpl.lib.wa.us/v2/nwroom/wanames.htm. 


CrisPIN S. Guppy, 4627 Quesnel-Hydraulic Road, 
Quesnel, BC V2] 6P8 Canada and Norbert G. 
KONDLA, Box 244, Genelle, British Columbia VOG 
1G0 Canada. 


Received for publication 5 November 2001; revised and accepted 
4 May 2002. 


BOOK REVIEWS 


Journal of the Lepidopterists’ Society 
56(3), 2002, 182-183 


BUTTERFLIES THROUGH BINOCULARS: THE WEST, A 
FIELD GUIDE TO THE BUTTERFLIES OF WESTERN 
NortH AMERICA, by Jeffrey Glassberg. 2001. Oxford 
University Press, Inc., 198 Madison Ave, New York, 
New York 10016. v—x + 373 pp., including 1076 color 
photos. Paperback, 8.25 x 5.5 inches, ISBN: 019- 
510669-5, $19.95. 


As organisms and their habitats succumb to the 
weight of encroaching humanity, we race to under- 
stand a little about our surroundings in hopes of con- 
serving what we have yet to destroy. We focus on 
groups with public appeal and use them to promote 
awareness of the negative impact that results from se- 
verely altering natural habitats. Butterflies have be- 
come the focus of some conservation inclined people, 
like birds before them. Jeffrey Glassberg’s Butterflies 
through Binoculars (BTB) series gives us a noncon- 
sumtive approach via the use of lenses, allowing for an 
up-close view of butterfly biology and identification 
without interfering with normal butterfly behavior— 
conservation, apparently, being the ultimate goal. 
BTB-The West, the latest in the BTB series, continues 


Glassberg’s view of butterflies and his photography of 


western North American butterfly fauna. 

The book has over 1100 photographs of butterflies 
(western United States and the southern part of west- 
ern Canada east to include all of Saskatchewan and the 
United States to just south of Houston), mostly in 
poses one would observe in the field. Accurate size 
relationships of species included on a page allow the 
user to compare size of similar looking species, and 
magnification or reduction from life size is indicated 
on each page. Some species have an inset of a different 
view of the species in question, while others have a 
black or white line to indicate a diagnostic feature. 
Photographs were arranged so that similar species are 
shown in similar poses, making comparisons easier. 
Dates and locations of the photographs are indicated 
in the rear of the book, providing a record of seasonal 
variation at particular localities. Species accounts that 
accompany photographs include habitat, abundance/ 
phenological, host plant, range, and other information. 
Color coded distribution maps indicate brood num- 
bers and species rarity or loss at a particular locality. 
Species and respective families are roughly organized 
phylogenetically, and common names, as recognized 
by the North American Butterfly Association’s 1995 
Checklist and English names of North American But- 
terflies are utilized throughout. 


The remainder of the book includes introductory 
sections on the use of binoculars and how and where 
to find butterflies (nectar sources, mud puddles, hill- 
tops, etc.) and how to identify them through binocu- 
lars, and a section on butterfly photography for those 
inexperienced behind the lens. Subsections under 
photography include information on equipment, photo 
etiquette, and caring for and viewing your pho- 
tographs. Brief sections on butterfly anatomy, biology, 
behavior and gardening are also included. Other intro- 
ductory sections include Glassberg’s thoughts on con- 
servation, netting butterflies, and commercially raised 
butterflies, and express his concern for these issues 
and the overall health of our butterfly fauna. A brief 
section on Hawaiian butterfly fauna, rare strays ob- 
served out west, and dubiously reported species can be 
found in the back of the guide. Supplementary text on 
species or groups that present identification problems 
(i.e., Euphilotes blues, Greater Fritillaries) are also in- 
cluded in the books rear and intended to ease identifi- 
cation difficulties in some regions of western United 
States. Foodplant scientific names, a list of major or- 
ganizations concerned with butterflies, a short glossary 
of butterfly terms, and a bibliography of western North 
American butterfly literature are also included. The di- 
mensions of the book are appropriate for a field guide, 
and print on pages holds up if moistened. 

This guide is written and intended for novice lepi- 
dopterists. Those seeking a complete list or pho- 
tographs of all species and subspecies in western 
North America, or a thorough taxonomic treatment 
will not find it here. The repeated use of the term “an- 
tennas” on page 10 (“antennas” are conductors by 
which electromagnetic waves are sent out or received 
and “antennae” are movable, sensory appendages oc- 
curring on the heads of insects and other arthropods) 
and “genuses” on pages 182 and 224 (genera was used 
on page 56) also give subtle hints to the intended audi- 
ence. 

It is admirable that Glassberg suggests an approach 
to butterfly observation, identification and conserva- 
tion with the production of the BTB series, a method 
that incidentally has been used for more than 50 years 
among tropical naturalists. His message, however, is 
over-biased to simple observation of butterflies 
through lenses for biology and identification purposes 
rather than legitimate, scientific research that is essen- 
tial to better understand these insects. There is already 
a mountain of bureaucratic nonsense that legitimate 
scientific research must contend with, a trend that will 
surely hinder rather than further our understanding of 
species relationships within environments. As the ma- 
jority of Glassberg’s audience will be novices, he 


VOLUME 56, NUMBER 3 


should not misinform with ideas that we can under- 
stand all butterfly biology and identification without 
research requiring sampling (yes, at times with a net) a 
small fraction of an overall species pool (very few col- 
lectors have used their nets to decimate the remaining 
populations of declining butterflies). In this regard, the 
book does not address the importance of conservation 
with a complete insight into the study of Lepidoptera. 
We absolutely cannot, and throughout history we 
could not have, come to the level of understanding (es- 
pecially the accurate identification) of butterflies by 
observing them through binoculars or photographs, 
and much remains to be discovered, even in North 
America. This book is for novice (perhaps naive?) lep- 
idopterists beginning a hobby, but would be of mar- 
ginal use to the experienced lepidopterist. 


JaMEs C. DuNFoRD, Department of Entomology 
and Nematology, University of Florida, Gainesville, 
Florida, 32611-0620, USA 


Journal of the Lepidopterists’ Society 
56(3), 2002, 183-184 


THE MILLENNIUM ATLAS OF BUTTERFLIES IN BRITAIN 
AND IRELAND, by J. Asher, M. Warren, R. Fox, P. 
Harding, G. Jeffcoate and S. Jeffcoate, Oxford 
University Press, 456 pp.; 270 color illus: publication 
2001; Cloth ISBN: 0-19-850565-5; Cloth Price: 
$40.00. 


The United Kingdom has had virtually no intact 
habitat for the past 500 years and it sports less than 
100 species of butterflies. Nonetheless, there are more 
books treating the butterflies of the British Isles than 
anywhere else on earth. So why do we need another 
book? Despite the area of coverage and topic this is 
not simply another butterfly book. It is a revelation. 
The six collaborative authors, whose professions in- 
clude physics, ecology, biochemistry and medicine 
have not produced something assembled by commit- 
tee. Rather, the fruits of their labor resulted in a de- 
cidedly readable book that is comprehensive, visually 
beautiful, scholarly and eminently valuable to a wide 
audience. The information content and style of the 
Millennium Atlas is a remarkable benchmark that fu- 
ture butterfly books must attempt to emulate. 

The backbone of the Millennium Atlas is the in- 
credibly comprehensive understanding of butterfly 
distributions. Collated by over 60 people in charge of 
coordinating local records, hundreds of thousands of 
observations were verified, entered into a central data- 


183 


base, plotted on a detailed geographic grid map whose 
resolution ranges from 10 km? to 100 m?, and then an- 
alyzed in the context of records spanning over 100 
years. Thousands of volunteers (amateur and profes- 
sional) contributed to what, in some cases, consist of 
many thousands of observations per species on each 
map. This is the template for the Millennium Atlas. 
The result is a magnificent understanding of the his- 
torical variation in distribution, population ecology, re- 
productive biology, colony dynamics, food plants, habi- 
tat use of each species, and a chronology of changes 
that reveal conservation successes, threats and failures. 

The detailed species accounts draw from the com- 
prehensive database and, of course, form the corpus of 
the book. Each account provides a full-page distribu- 
tion map, a color photograph of the butterfly taken in 
the wild, and a complete account of food plants, habi- 
tat, lifecycle and colony structure, local distributional 
trends contrasted with those on the European main- 
land, a summary of the ecological and conservation 
outlook for each species, and references to recent lit- 
erature. Each species account is a pleasure to read, 
and a work of art into the bargain. Therefore the book 
will be of use to virtually anyone with an interest in 
butterflies and their place in nature. 

Tucked away at the end are nine appendices ranging 
in content from vernacular names and addresses 
where contributing data should be sent, to lucid 
graphs showing dynamical changes in the abundance 
and distribution of many species. These are followed 
by a comprehensive bibliography that provides the 
reader with a tremendous amount of research possibil- 
ity. Calling these sections thorough hardly gets into the 
ballpark, or if you prefer, the cricket pitch. These sec- 
tions are vital examples of how scholarly information 
can be presented in admirable and easily accessible 
form. 

This volume resonates a cultural history that em- 
phasizes cognizance of the natural world. For genera- 
tions, denizens of the British Isles have shown an abid- 
ing interest in natural history, a proclivity to make 
abundant and accurate observations, share them, and 
interpret those observations for understanding wildlife 
and its conservation. Such a historical cultural interest 
in butterflies has fostered, and ultimately led to, the 
Millennium Atlas. However, the Millennium Atlas is 
not a finished work that heralds the end of butterfly 
study in Britain. Rather, the recent collaborative ef- 
forts of many European biologists working on butter- 
flies in a fragmented landscape attests to it being a 
springboard to further refine our dynamic range of un- 
derstanding butterflies. In producing the Millennium 
Atlas the cultural heirs of Darwinism elegantly demon- 


184 


strate the evolution of butterfly populations in the con- 
text of a changing environment. Those of us who work 
in areas where so little is known about butterflies may 
be daunted by such an effort. Nevertheless, it is clear 
we must strive toward its example. 

Sturdily bound, filled with illustrations and informa- 
tion, and inexpensive, this book is uniquely outstand- 
ing in many ways. However, I especially appreciated 
one aspect of it; the subtle yet pervasive conservation 
underpinning based on the study of butterflies. If I 
read the message correctly it might be paraphrased in 
a more general way: appreciate and learn from nature 
before it vanishes into the increasingly greedy maw of 
the human dominated landscape. That is to say, the 
game is not just about collecting and writing epitaphs 
of colorful insects. It is about preventing the ecological 
holocausts that surround us, and surviving. As the song 
title suggests, now is the time. 


P. J. DEVRIES, Center for Biodiversity Studies, 
Milwaukee Public Museum, 800 West Wells St., 
Milwaukee, Wisconsin 53233, USA 


Journal of the Lepidopterists’ Society 
56(3), 2002, 184-188 


BUTTERFLIES THROUGH BINOCULARS: THE East. A 
FIELD GUIDE TO THE BUTTERFLIES OF EASTERN 
NortH AMERICA, by J. Glassberg. 1999. Oxford 
University Press. $18.95. ISBN 0-19-510668-7 


HOW MUCH SCIENCE IS TOO LITTLE? 


The practice of butterfly watching has rocketed in 
recent years, fueled primarily by increasingly accessi- 
ble field guides equipped with easy shortcuts to identi- 
fication, technological advances in photography, and 
numerous organizations, local and national, devoted to 
furthering public awareness of butterflies and their di- 
versity. Butterfly watching, a healthy medium for natural 
history education, has realized considerable popularity 
not just among weekend naturalists and recovering bird- 
watchers, but also among educators and even profes- 
sional lepidopterists that participate in local butterfly 
counts. Dr. Glassberg’s Butterflies through Binoculars: 
The East (hereafter BTB) is likely to surpass previous 
guides’ popularity among butterfly watching enthusi- 
asts, and to that end it serves as a photographic guide 
for most eastern butterflies. Unfortunately, the general 
utility of this book is reduced sharply by several short- 
comings, all of them related to the furtherance of lep- 
idopteran science and science-based conservation. 

It is difficult to decide in what spirit to review this 
book. It is clearly a volume intended for hobbyists 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


(which is fine), but at the same time the purview of its 
influence is intended to include matters of scientific 
import, and it includes promotional material linked to 
the North American Butterfly Association (NABA). 
The historical and organizational context from which 
the butterfly-watching movement has sprung is plain 
within the pages of BTB, and I therefore view it as im- 
possible to review this book and some of the informa- 
tion presented therein independently of similar mes- 
sages published by NABA. It is my hope that a 
contextual review of this kind will add more light than 
heat. Regardless, after tending to the book's technical 
aspects, I will proceed to its message. 

First to the nitty gritty. The meat of BTB is of course 
its 283 species accounts and accompanying pho- 
tographs. Again, through these, BTB is a fine introduc- 
tion to observing butterflies of eastern North America. 
Species accounts include identification cues, geo- 
graphic range maps (which accompany the plates), 
some life history information, including primary host 
plants and extensive flight season data in the form of 
“phenograms” which consist of rough relative abun- 
dance by month in four geographically separated states 
(WI, NY, NC, and LA) generated in consultation with 
various local experts (no use of collection data is refer- 
enced). The lack of detailed descriptions should not be 
viewed as a shortcoming, assuming the guide’s primary 
target audience comprises hobbyists and prospective 
inventorists. The identification cues are by and large 
well crafted, with the exception of their reliance on 
comparative observation: Size measurements for 
species are not generally given, but evaluated with ref- 
erence to other butterflies. Field diagnoses, when 
present, are emboldened, and brief descriptions ac- 
companying plates make quick identification easy and 
reduce the need for page flipping. The photographic 
quality is generally quite good (with a few exceptions), 
and should help butterfly watchers get a feel for what 
various species look like in vivo, although field marks 
are not consistently delineated. The photographs are 
also carefully scaled against others on the plate. In any 
event, BTB should serve as a step towards identifying 
butterflies reliably in the field, and therefore meets its 
purpose. 

The text comprising the book’s introduction in- 
cludes sections on butterfly photography, butterfly gar- 
dening, tips on finding butterflies, and butterfly biol- 
ogy. There is no discussion of proper vouchering, 
collecting, or rearing techniques or protocols, nor any 
broad discussion of butterfly taxonomy or systematics 
except for some remarks buried in the species ac- 
counts. With those exceptions, this material serves as 
an adequate introduction for the casual butterflyer. 


VOLUME 56, NUMBER 3 


Text following the species accounts includes a bibliog- 
raphy and a short glossary of terms. 

I detect three principal shortcomings in the book’s 
message, all associated with teaching science-based 
natural history and the relevance of that science to 
conservation of landscapes and species. First, Dr. 
Glassberg’s dismissal of the nomenclatural process 
combined with the assertion that butterfly taxonomy, 
including both scientific and common names, is being 
“standardized” (p. 33) by NABA is a serious shortcom- 
ing. It also may render BTB impotent in serious bio- 
inventories, butterfly counts, or identifying popula- 
tions of cryptic species. Second, while no doubt well 
meaning, Dr. Glassberg makes a number of misleading 
comments about managing butterfly habitat, specifi- 
cally fire management, that may oversimplify and ex- 
acerbate an already controversial conservation issue. 
Third, BTB’s strident indictments of collecting utterly 
fail to mention the critical importance of collections 
not only to the study of butterflies, but also the fur- 
therance of taxonomy, systematics, and the conserva- 
tion and protection of threatened species. 

There have always been nomenclatural controver- 
sies in the butterfly literature, for that is the nature of 
science: To modify existing taxonomy as new entities 
are discovered and described and as recent informa- 
tion is brought to bear on our imperfect understanding 
of nature. The proliferation of names is perhaps par- 
ticularly acute in the most showy organisms, including 
birds, butterflies, and tiger beetles, because the atten- 
tion they draw from hobbyists leads inevitably to the 
discovery of novel forms that are subsequently de- 
scribed and all too often given subspecific or infrasub- 
specific epithets. To be sure, keeping up with current 
nomenclature by tracking the most recent revisions 
and extracting valid names is a difficult task, one 
deemed necessary for scholars but annoying to those 
with a more casual interest. To be sure, the existence 
of privately published, non-peer-reviewed journals 
that never make their way to libraries but in which 
new species and subspecies are regularly described is 
a bane to taxonomy, and frustrating to anyone at- 
tempting to incorporate current nomenclature into 
field guides. It is thus not uncommon for authors to 
prefer, explicitly or otherwise, particular taxonomic 
arrangements on an ad hoc basis. The nomenclature in 
BTB, both scientific and vernacular, follows NABA’s 
Checklist and English Names of North American But- 
terflies. And while there are some differences of opin- 
ion and treatment of taxonomic status are evident be- 
tween BTB and other field guides, for example, the 
most pointed remarks in BTB are directed at the 
nomenclatural process itself. Obviously dissatisfied 


with the state of flux in North American butterfly 
nomenclature, Dr. Glassberg writes (p. 33): 


“Before the NABA checklist was published in 1995, 
each author of a book about butterflies used whatever 
set of names struck his or her fancy. The result has 
been a confusing plethora of names that has bewil- 
dered the uninitiated and made it more difficult for the 
public to become involved with butterflies. We are now 
on the road toward standardization, although this 
process will take years to be completed.” 


Later Dr. Glassberg writes (p. 53) in the context of 
the species account for the Mustard White Pieris napi: 


“There is some recent evidence that the Mustard 
White complex may consist of a number of different 
species and a number of books have appeared that 
jump on this bandwagon. Changes in well established 
names should be made in works intended for the pub- 
lic only if the published evidence is overwhelming 
[emboldening as in BTB] that the change is correct.” 


Notwithstanding that fact that, like many other strong 
opinions expressed in BTB, these statements are unac- 
companied by specific references, these assertions also 
belie either a fundamental misunderstanding or a disre- 
gard for the scientific process and its critical value to bi- 
ological conservation. As most lepidopterists know, there 
are a number of competing endeavors to “standardize” 
butterfly taxonomy. Difficult as it may be for those 
with a superficial interest in butterflies to keep up with 
the technical literature, are we to believe that taxonomic 
stability, via the acceptance of one person's sanctioned 
list is more important than taxonomic progress via 
scholarly study? Dr. Glassberg seems to find taxonomy 
and systematics so trivial as to make them beholden to 
the hobbyist. In certain cases (e.g., p. 153), taxonomic 
progress is explicitly ignored and excused only in order 
to remain consistent with the NABA checklist. 

Unfortunately, the problem goes further: NABA 
publishes the results of annual butterfly counts, but 
with little quality control. Thus it is easy for erroneous 
records to proliferate in print, and such apparent 
“data” could, if taken seriously, prove fatal to distribu- 
tional revisions and local conservation efforts. For ex- 
ample, I recently encountered a local butterfly check- 
list, compiled by an avid (and talented) butterfly 
watcher, encouraged by a prominent international con- 
servation organization, and copyrighted. The checklist 
consisted primarily of records previously published by 
lepidopterists (who were not credited) and a number 
of new records for the region in question for which no 
specimen vouchers exist. Some of these species were 
described as occurring commonly. It might be worth- 


186 


while to contemplate the impact of such information 
should these species ever fall in need of protection. 
Consider, for example, the unnecessary obstacle to se- 
curing protection of a species presented when indica- 
tions that it is abundant are disseminated. Such scenar- 
ios are only worsened when taxonomic progress is 
ignored in favor of a popular “standard.” Bottom line: 
The combination of inflexible and possibly question- 
able standardization with an unverifiable system of 
record keeping renders any inventories or butterfly 
counts meaningless without substantial follow up. 
“Standardization” won't change the valid names, and if 
butterfly watchers promote ignoring those, then their 
records will be even more meaningless. 

Dr. Glassberg’s own anti-collecting agenda, and that 
of his organization, NABA, is well known. But as with 
a number of issues on which Dr. Glassberg purports to 
speak authoritatively, intelligent discussion of this 
complicated and controversial issue is done a disser- 
vice by the author's treatment in BTB and elsewhere. 
As a lifelong conservationist and lepidopterist, I have 
been impressed by the unique ability of the lepi- 
dopterist community to integrate professional biolo- 
gists with so-called “amateurs.” I think this is recog- 
nized by most as one of the major strengths of 
lepidopteran research. True, controversies have arisen 
with respect to such issues as collecting regulations 
and the listing of endangered species, and irresponsi- 
ble collectors do indeed exist. But in my experience 
those lepidopterists most effective at resolving such is- 
sues have done so by thinking clearly, speaking articu- 
lately, collecting hard data, writing with skill and schol- 
arship, and all without resorting to divisive tirades. 
Alas, such cannot be said of BTB. 

At times, Dr. Glassberg’s stance on collecting (and 
nomenclatural standards) appears predicated on the 
notion that our understanding of the North American 
fauna is sufficient, if not complete. One could, per- 
haps, make such a case for birds, and butterflies’ being 
the birds of the insect world; the temptation to think 
likewise for them is perhaps understandable—at least 
for a birdwatcher. But think for a moment on the real 
state of lepidopteran taxonomy. Do we really have a 
complete enough understanding of butterfly variation, 
even in the conterminous United States, to stop study- 
ing it? I know of numerous recently discovered or un- 
described North American species. Lepidoptera are 
notorious for sudden range changes, both contractions 
and expansions, as well as for host plant shifts and 
rapid speciation events. Regulators and legislators do 
not frequently distinguish recreational from scientific 
collecting, and if scientific collecting were removed 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


from the equation, the endeavors of scholarly research 
would be hamstrung. The question is whether serious 
scientific research should take a back seat to the per- 
sonal views, however noble, of those who don’t under- 
stand the importance of collecting. 

But the importance of collections is certainly not 
limited to the realm of systematics. Our understanding 
of conservation priorities would not exist without the 
information contained in zoology collections and 
herbaria. Importantly, our ability to protect popula- 
tions and use the occurrences of rare organisms to pre- 
vent destructive development and land use practices 
often depends on voucher specimens’ being deposited 
in scientific collections. I have known would-be devel- 
opers to contest the occurrence of rare Lepidoptera on 
their properties, often during conservation commis- 
sion hearings in mid-winter when vouchering is im- 
possible, and were it not for voucher specimens in col- 
lections, important natural areas would have suffered. 

It seems to me that Dr. Glassberg is too busy prose- 
lytizing to present a balanced discussion, or to bother 
mentioning the importance of scientific collections. In- 
deed, he appears to go to considerable lengths to 
downplay the potential role of amateurs in contribut- 
ing to them. The recent Dragonflies through Binocu- 
lars field guide, published under the auspices of 
NABA, contains not an iota of information on how to 
properly prepare dragonfly and damselfly specimens. 
Here is a group of organisms about which we know 
significantly less distributional information than but- 
terflies, presenting an outstanding opportunity for am- 
ateur naturalists to make valuable contributions to sci- 
ence and conservation. For someone so concerned 
about the public’s access to information, Dr. Glassberg 
seems content to limit it selectively. I can't wait to see 
Moths through Binoculars. 

Assuming trumping nomenclatural practices will 
amount to nothing, perhaps the most disturbing and 
disingenuous aspect of Dr. Glassberg’s agenda is his 
stance on conservation. He describes (p. 27) conserva- 
tion as the “raison d’etre of this book” and to the extent 
that it will help instill an appreciation of the natural 
world and enhance access to its study, it may make a 
contribution. Unfortunately, both BTB’s message and 
its intended furtherance of conservation are hampered 
by a series of misleading comments (as well as by the 
adherence to a rogue taxonomy). Among the more 
disingenuous are those directed at the use of pre- 
scribed burning in managing natural areas. In Dr. 
Glassberg’s (p. 28) words, fire is an “often misguided 
conservation tool,” and goes on to smear the conserva- 
tion community (p. 29): 


VOLUME 56, NUMBER 3 


“Unfortunately, some of the conservation commu- 
nity and groups charged with the management of our 
natural areas have made a devil's embrace of fire—‘“fire 
is good at keeping areas open, let's burn like crazy.” 


Dr. Glassberg only begrudgingly acknowledges the 
importance of responsible fire management, but his 
overall message is to alert naive readers to what he 
characterizes as widespread irresponsibility in the ap- 
plication of prescribed burning by land managers. 
Granted, we are all aware of examples of poor fire 
management, much as we know of examples of irre- 
sponsible politics, business practices and, yes, collect- 
ing. But the fact remains that only a small percentage 
of North America’s fire dependent communities are 
currently being managed with fire at all. 

Later, he writes (p. 30): 


“[A] conservation professional, trained to look for 
“pure” examples of native habitats, may take one look 
at these weedy fields [containing important nectar 
sources] and turn up her/his nose.” 


One is left to wonder whose training and credentials in 
entomology or conservation are worthy of Dr. Glass- 
berg’s approbation. 

It is interesting to observe that BTB’s crusade 
against science and science-based conservation has 
been extended in an indictment of how conservation 
agencies operate. Using NABA as a vehicle, Dr. Glass- 
berg has recently called upon The Nature Conser- 
vancy and the Heritage Program network to release in- 
formation on locations of threatened plant and animal 
occurrences, arguing that the public should have un- 
limited access to such information (Glassberg, 2001). 
In a recent editorial of American Butterflies Glassberg 
(2001:2), writes: 


“The Nature Conservancy, other conservation organ- 
izations, and government agencies charged with protect- 
ing our flora and fauna, keep the locations of rare plants 
and animals a secret with the best of intentions: obsessive 
collectors are a threat—especially to certain populations 
of rare orchids, cactuses [sic], butterflies, and reptiles.” 


This is an interesting, but inaccurate and incomplete 
characterization of why threatened species informa- 
tion is kept confidential. In fact, most of the reasons 
have nothing whatsoever to do with collectors, but 
rather with protecting the rights of landowners and 
protecting the most vulnerable sites from habitat de- 
struction. As anyone with experience in serious bioin- 
ventories and conservation assessment knows, many 
private landowners would be loathe to cooperate with 
conservation endeavors if anyone could access rare 


187 


species information for their properties. In my experi- 
ence, protecting landowner privacy has been critical to 
conserving some of North America’s most significant 
natural areas. Equally important is the issue of illegal 
habitat destruction. What is to prevent a would-be de- 
veloper of a shopping center in a wetland, for example, 
of bulldozing the site at midnight upon finding out it 
contains a colony of some rare orchid. No rare butter- 
fly or orchid, no hassles with Fish and Wildlife. It may 
sound trite, but it happens, and clearly tightening col- 
lecting regulations is not a solution. Serious scholarly 
study is. 

Unfortunately, rather than contribute to disseminat- 
ing useful information and understanding to the pub- 
lic, it appears that Dr. Glassberg prefers an alarmist 
route. In the same editorial Glassberg (2001:2) writes: 


“Butterflies through binoculars: The East describes 
how collectors killed the last Mitchell’s Satyrs in New 
Jersey, invading private property despite the fact that 
the owner, in a valiant attempt to save the butterflies, 
encircled the land with chain link fences and posted 


guard dogs.” 


But perhaps the most telling quote from the editorial 
is this: 


“Last summer, NABA petitioned the U.S. Fish and 
Wildlife Service to list Miami Blues as federally endan- 
gered on an emergency basis. We decided to keep the 
location of the colony secret until listing could provide 
some protection. While I understand that the petition 
was favorable [sic] received, the Department of Inte- 
rior subsequently issued a moratorium on all listings 
and it is unclear when, if ever, this species will be pro- 
tected. So, I have decided to make the location of the 
colony public {my italics added]. I hope that Secretary 
of the Interior Norton acts to list this species and that 
Florida legislators enact laws that provide real protec- 
tion, before it too late, but I am not optimistic. So, my 
advice to you is to see these butterflies, at the northern 
end of Bahia Honda State Park, along Silver Palms Na- 
ture Trail, while you still can. But please don't tell any- 
one else.” 


Now let us get this straight: Even given that the site 
occurs in a state park, presumably protected from de- 
velopment, Dr. Glassberg divulged specific site infor- 
mation in virtually the same breath as noting that the 
maximum fine is $50 and that prospects for serious 
protection are unlikely. Even if illegal collecting were 
a threat (which for all I know it might be in this case), 
hasn't Dr. Glassberg just declared open season on this 


butterfly? 


188 


To many it is unfortunate that Dr. Glassberg 
chooses to treat complex and controversial scientific is- 
sues that bear on the discovery and understanding of 
nature by resorting to misinformation and spurious ap- 
peals from the safety of his editorial fiefdom. But by al- 
lowing the propaganda to spill into BTB, he corrupts a 
potentially useful book with an agenda-driven crusade 
against science and scientific conservation. As such, 
much of BTB’s utility is lost through the use of rene- 
gade nomenclature and idiosyncratic presentation of 
important issues. 

In summary, BTB is valuable as an introduction to ob- 
serving and photographing butterflies, but its failure to 
deal responsibly with serious conservation-related and 
scientific issues can, in my opinion, only result in further 
muddying waters in dire need of clearer solutions. 


LITERATURE CITED 


GLASSBERG, J. 2001. Listen, do you want to know a secre (Do you 
promise not to tell) [sic]. American Butterflies 9(3):2 9. [Editorial] 


PAUL GOLDSTEIN, Division of Insects, Field Museum 
of Natural History, Roosevelt Road at Lake Shore 
Drive, Chicago, Illinois 60605-2496, USA 


Journal of the Lepidopterists’ Society 
56(3), 2002, 188-189 


CATALOGUE OF THE GENERA OENEIS AND DAVIDINA, by 
Vladimir Lukhtanov and Ulf Eitschberger. Part 11 and 
Supplement 4 of E. Bauer and T. F iaakenbachs eds., 
BUTTERFLIES OF THE WORLD, Goecke and Evers, Kel- 
tern, Germany, 2000 (Part 11, Plates) and 2001 (Sup- 
plement 4, Text). Plates (28 plates + 12pp; ISBN 3- 
931374-81-5) and text (37pp.; ISBN 3-931374-52-1) 
available from Antiquariat Goecke & Evers, Thn. Erich 
Bauer, Sportplatzweg 5, 75210 Keltern, Germany 
(www.insecta.de). Prices Euro 22.50 (plates) and 20.00 
(text). 


This work, Part 11 of the ongoing Butterflies of the 
World series edited by Bauer and Frankenbach, is a 
complete illustrated catalog of the genera Oeneis and 
Davidina (Lepidoptera: Nymphalidae, Satyrinae, 
Oeneini). The Plates volume comprises 28 excellent 
color pages that show, at life size, examples of each of 
the 199 taxa catalogued. In general, both sexes are il- 
lustrated with the upper- and under-sides of each spec- 
imen shown on facing plates. This volume includes 
complete data (cited verbatim from the specimen la- 
bels) and brief discussions of the distribution of each 
taxon. Thirty-five of the approximately 350 individual 
specimens illustrated are primary types. Incidentally, 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


the German-language edition of Part 11 (ISBN 1- 
931374-80-7) was published before the English version 
and includes the original descriptions of Oeneis buddha 
greishuberi, O. b. frankenbachi, and O. tarpeja baueri. 
These descriptions are not in the English version. 

The Text, published as Supplement 4 to Part 11, in- 
cludes a key to the 10 species-groups recognized by 
the authors and a discussion of each species and sub- 
species. Full citations to the original descriptions, 
complete synonymies, locations of type material, and 
excellent-quality distribution maps are provided. Male 
and female genitalia are shown for several taxa in each 
species-group. The English text includes description 
of one new subspecies (Oeneis nanna taimyrica), ab- 
sent from the German edition. In addition, three neo- 
or lectotypes are designated and 3 new combinations 
are established, and 1 species and 2 subspecies names 
are reduced to synonymy. The tribe Davidini Cho Io 
1998 is synonymized with Oeneini Wheeler 1903. 

It is a lovely thing to have before one’s eyes a com- 
plete holarctic genus, males and females, dorsal and 
ventral. This is especially true in the case of a genus 
whose speciation has been extensive in the mountains 
of central Asia and China, where the existence of such 
outstanding species as mongolica, urda, and buddha is 
unsuspected by perhaps most North American lepi- 
dopterists. Likewise, it is a revelation to see specimens 
of jutta ssp. from localities ranging from Colorado to the 
Altai mountains on the western border of Mongolia to 
Siberia. And the two strange, pierid-like Chinese species 
of Davidina (with genitalia very close to Oeneis but dis- 
tinct venation) are like nothing else among the satyrs. 

Oeneis is a difficult genus and the authors are care- 
ful to point out a number of areas where further re- 
search is needed. For instance, lucilla Barnes and Mc- 
Donnough 1918 is retained as a melissa ssp, but full 
species rank is “not excluded”. It is pointed out that 
adult morphology is insufficient to resolve difficulties 
in taxonomy of the O. norna group (norna, rosovi, 
polixenes, philipi, actaeoides) and that study of DNA 
and immature stages is needed. O. ivallda (Mead 
1878) is retained but its conspecifity with chryxus 
“cannot be excluded”. 

This work was made possible largely through assem- 
blage, in the Entomologische Museum Eitschberger, 
of a worldwide collection of 4400 specimens of Oeneis. 
Through study of this material (supplemented by re- 
search in the Zoologische Staatsammlung in Munich 
and the Zoological Institute in St. Petersburg, Russia, 
and study of material from the private collections of 
Grieshuber, Zhou, Strohle, Tremblay, Ferris, and oth- 
ers), Lukhtanov and Eitschberger have produced a 
work that combines solid scholarship and elegant pre- 


VOLUME 56, NUMBER 3 


sentation. The reasonably priced work should be on 
the shelf of any lepidopterist with an interest in mon- 
tane and arctic Satyrinae. 


Joun A. Hyatt, 5336 Foxfire Place, Kingsport, 
Tennessee 37664, USA, e-mail jhyatt@eastman.com 


Journal of the Lepidopterists’ Society 
56(3), 2002, 189 


THE MOTHS AND BUTTERFLIES OF GREAT BRITAIN AND 
IRELAND, vol. 4, edited by A. M. Emmet and J. R. 
Langmaid. 2002. Published by Harley Books, Martins, 
Great Horkesley, Colchester, Essex CO6 4AH, England. 
Part 1 [Oecophoridae, Ethmiidae, Autostichidae, Blasto- 
basidae, Batrachedridae, Agonoxenidae, Momphidae, 
Cosmopterigidae, Scythrididae]: 326 pp., 95 text figures, 
146 maps, 13 color plates; Part 2 [Gelechiidae]: 277 
pages, 63 text figures, 161 maps, 6 color plates. Hard- 
back, 26.2 x 20.8 cm, ISBN 0 946589 63 1 [set], ISBN 0 
946589 66 6 [part 1], ISBN 0 946589 67 4 [part 2]. Price 
£80.00 [pounds sterling] per part, $150.00 jacketed set 
excluding postage, available from publisher. 


This two-part volume is beautifully produced: the 
typography is clear; the line drawings of genitalia, 
heads, venation, and larval activity are extremely well 
done and accurate; and the color plates are outstand- 
ing. Second, it represents a clear, concise guide to the 
gelechioid fauna [exclusive of Coleophorinae and 
Elachistinae treated in volume 3] of Great Britain and 
Ireland. This work is not, and does not pretend to be, 
revisionary. Nomenclature closely follows the most re- 
cent comprehensive list of taxa presented by John 
Bradley (Log book of British Lepidoptera, 2000). Thir- 
teen authors contributed the nine families (147 species) 
in part 1, 10 to the Gelechiidae (160 species) in part 2. 

In my early years as a budding systematist the only 
line drawings of microlepidopteran genital characters 
were in The genitalia of the tineid families of the Lepi- 
doptera of the British Islands by Pierce and Metcalf 
(1935). They were relatively crude and did not well il- 
lustrate many important features; however, they 
formed the basis of my knowledge of the western Eu- 
ropean gelechioid fauna for many years. With the pub- 
lication of volume 4 of The Moths and Butterflies of 
Great Britain and Ireland this body of information has 
been advanced significantly 

Family-group treatments comprise a statement on 
geographic distribution, classification, morphologic 
characters of adults and immatures, larval hosts and 
habits, pupation sites, phenology, adult habits, a check- 
list, identification key to genera, illustrations of male 


189 


and female genitalia, and references. Particularly use- 
ful is the illustrated key to the genera of Gelechiidae 
based on male genital characters. All genera and 
species are characterized following the pattern for 
family-groups but with the addition of the synonymy 
and literature citation for each name, illustration of ve- 
nation for each genus, lateral views of selected heads, 
a distribution map for each species, larval hosts and 
habits, and adult habits and summary of geographic 
distribution. The historical record of each species pro- 
vides insights to faunal changes. Specific variation is 
noted, and contrasts with similar species are clear. 
Much valuable information, which should prove useful 
for other temperate taxa, is contained for each taxon. 

The quantity of life history information is impres- 
sive. Beyond the relatively standard discussion of larval 
appearance and host plants are treatments of larval, 
pupal, and adult behavior; habitats where adults may 
be found; diel activity of adults; and history of occur- 
rence. The latter reveals interesting and useful infor- 
mation such as: 1) species known only from Great 
Britain and collected once, Euclemensia woodiella 
(Curtis); 2) species collected once in Great Britain but 
naturally occurring elsewhere, Epicallima formosella 
([Denis & Schiffermiiller]); 3) species known from a 
single locality and regularly collected (Monochroa 
niphognatha (Gozmany), Metznaria littorella (Doug- 
las)); 4) species very infrequently collected (Dichome- 
ris ustalella (F.) collected in 1861, 1987, and 1999): 5) 
species with general occurrence (Agonopterix nervosa 
(Haworth) and Endrosis sarcitrella (L.)): and 6) intro- 
ductions (Scythris inspersella (Hiibner)). Because col- 
lection data have been, and continue to be, collected 
and databased, and detailed knowledge of larval hosts 
has accumulated, it is possible that realistic statements 
about abundance and distribution of gelechioid 
species can be made for the region and remedies pro- 
posed for maintenance of restricted populations. 

An anomalous feature of the entire series is a set of 
essays/discussions on topics related to Lepidoptera. In 
volume 4(1) (pp. 11-41, col. pl. A-F, fig. 1-6, tab. 1-3) 
J. Rydell and M. R. Young have an excellent review 
“The Ecology and evolution of Lepidopteran defences 
against bats,” which deserves more broad readership 
than is likely when the title appears solely in the table 
of contents. 

This volume sets a very high standard for compara- 
ble works on the fauna of individual countries. I con- 
gratulate all involved in the text, illustration, and pro- 
duction for an excellent product. 


RONALD W. HODGES, 85253 Ridgetop Drive, Eu- 
gene, Oregon, 97405-9535 USA 


ERRATA 


Journal of the Lepidopterists’ Society 
56(3), 2002, 191-192 


THE LARGE MOTHS OF GUANA ISLAND, BRITISH VIRGIN ISLANDS: A SURVEY OF EFFICIENT 
COLONIZERS (SPHINGIDAE, NOTODONTIDAE, NOCTUIDAE, ARCTIIDAE, GEOMETRIDAE, 
HYBLAEIDAE, COSSIDAE) 


In the above paper by Vitor O. Becker and Scott E. Miller (Journal of the Lepidopterists’ Society 56(1):9-44), 
specimens that compose a plate were numbered incorrectly. The corrected figure numbers and accompanying leg- 
end are given here. 


192 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 112-150. Arctiidae (112-124), Geometridae (125-146), Cossidae (147-149) and Hyblaeidae (150) (specimens from Guana, unless 
stated otherwise). 112, Hypercompe simplex, male (Puerto Rico); 113, Composia credula, male; 114, H. simplex, female; 115, Calidota strigosa, 
male; 116, Eupseudosoma involutum, male (Puerto Rico); 117, Utetheisa ornatrix, male; 118, U. pulchella, female (Brazil); 119, Empyreuma pu- 
gione, male; 120, Horama panthalon, male; 121, H. pretus, male; 122, Cosmosoma achemon, male (St. Thomas); 123, Eunomia colombina, male; 
124, Nyridela chalciope, female (Cuba); 125, Pero rectisectaria, male; 126, P. rectisectaria female; 127, Oxydia vesulia, male; 128, Erastria de- 
crepitaria, male (Cuba); 129, E. decrepitaria, female; 130, Sphacelodes fusilineatus, male; 131 S. fusilineatus, female; 132, Macaria paleolata, 
male; 133, Patalene ephyrata, male; 134, Almodes terraria, male (Bahamas); 135, Semaeopus malefidarius, male; 136, Leptostales noctuata, 
male; 137, L. noctuata, female; 138, Obila praecurraria, female (Tortola); 139, P. defensata, male; 140, P. defensata, female; 141, Eueana sim- 
plaria male; 142, E. simplaria female; 143, Phrudocentra centrifugarium, male; 144, P. centrifugarium, female (Cuba); 145, 146, P. centrifugar- 
ium, females; 147-149, Psychonotua personalis, males; 150, Hyblaea puera, male (Cuba). 


EDITORIAL STAFF OF THE JOURNAL 


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Associate Editors: 
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NOTICE TO CONTRIBUTORS 


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Palate: cee ee K ey es wean nena 


A REVISION OF THE GENUS BuLiA WALKER (LEPIpopTERA: Nocrupat) ; 
. A. Carolina Laughlin ic ne epeenis Tar as 


Francis H, Snow's New Mexico BUTTERFLY COLLECTIONS Cee Je 


DisTRIBUTION OF THE DIANA F RITILLARY, SPEYERIA DIANA (NyMPHALIDAR) 
NECTAR PLANT AND HABITAT PREFERENCE Matthew D. Moran and 


Genenat Nores 


| Nose CORRECTNESS OF Pacer PRATENSIS VS. i peecoe PULCI 
Norbert G. Kondla and Crispin S. Guppy ee 


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© This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanance of Paper). 


Na Ee. 
eles 
Los | 
Rh ase WA . f ae ; ela gee Volume 56 Number 4 
a= N aes Kins ; 9 December 2002 
il Sets 2 ISSN 0024-0966 


— Journal of the 
| lLepidopterists Society 


THE LEPIDOPTERISTS’ SOCIETY 

Executive Counci. at oe 

Lawrence F. Gatu, President Scorr E. Miiter, Vice President fe 
J. Donatp LaFonTainE, Immediate Past President Konrap Fiepier, Vice President 


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Ernest H. Wituiams, Secretary 


Members at large: 


J. Bolling Sullivan James K. Adams William E. Conner 
Philip DeVries Wayne F. Wheling Rebecca Simmons 
David Ahrenholz Jeffery R. Slotten Charles V. Covell Jr. 


EprroriaL BoarD 


Rosert K. Rossins (Chairman) ‘aor aes, Re 
Caria M. Penz (Journal) APS AY : : 
Wituiam E. Miter (Memoirs) + Cet aN rig 
Puiu J. Scuarrert (News) ae , Soe hae \ 

Joun A. Snyper (Web) ? Ren orn ah 


Honorary Lire MEMBERS OF THE SOCIETY 


Cxarves L. Remincron (1966), E. G. Munror (1973), IAN F. B. Common (1987), . ne Serene 
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Cover illustration: Detail of the forewing ventral surface of Morpho achilles (Nymphalidae): 


JOURNAL OF 


Tue Lepiporprerists’ SOCIETY 


Volume 56 


Journal of the Lepidopterists’ Society 
56(4), 2002, 193-198 


2002 


Number 4 


EXPERIENCE-RELATED CHANGES IN THE BRAIN OF 
AGRAULIS VANILLAE (L.) (NYMPHALIDAE) 


VADIM KROUTOV 
123 Bartram Hall, Department of Zoology, University of Florida, Gainesville, Florida 32611, USA 


ROGER L. REEP 
College of Veterinary Medicine, Health Science Center, P.O. Box 100144 Gainesville, Florida 32610-0144, USA 


AND 


TOK FUKUDA 


Center for Medical, Agricultural and Veterinary Entomology, Agricultural Research Service, 
U.S. Department of Agriculture, Gainesville, Florida 32604, USA 


ABSTRACT. 


In the brain of Agraulis vanillae, the size of the brain regions involved in the processing of olfactory information was found 


to depend on the butterfly’s experience. Butterflies collected in nature have olfactory glomeruli and mushroom body calyxes of larger relative 
size than do butterflies reared and kept in the laboratory in isolation from normal environmental stimuli. No size difference was found in the 


optic lobes or the central body in either males or females. 


Additional key words: 


The brain of an insect is the principal associative 
center of the body. It receives sensory information 
from a variety of sense organs, processes it and con- 
trols all functions of the organism, including complex 
forms of behavior. Several regions of the brain differ- 
ing in morphology and function are recognized and re- 
ferred to as neuropils (Fig. 1). Neuropils are the cen- 
ters of the regions and are formed by a complex of 
densely packed nerve fibers. The neurons, which com- 
pose a region, lie at its periphery. On histological sec- 
tions of the brain neuropils appear as much denser, 
darker than the rest of the brain areas. 

The neuropils of particular significance in the pro- 
cessing of information in insect brains are the mush- 
room bodies and antennal lobes. Mushroom bodies 
receive signals from different sense organs and experi- 
ments on Drosophila (Heisenberg et al. 1985, Han et 
al. 1992) and Apis (Erber et al. 1980, Menzel et al. 
1974, Hammer & Menzel 1998) suggest that they are 
implicated in olfactory memory formation. They are 
composed of three types of cells: cells that direct sig- 


mushroom body, neuropil, olfactory lobes. 


nals to the mushroom bodies, cells that deliver signals 
from the mushroom bodies to other parts of the ner- 
vous system, and the intrinsic cells (Kenyon cells) that 
connect the first two types between themselves. The 
Kenyon cells occupy the area around the mushroom 
body neuropil. 

All information from the organs of smell (olfactory 
organs) is received in another brain region: antennal 
lobes, which are critically important in the delivery of 
olfactory information to the mushroom bodies. An- 
tennal lobes are composed of a series of neuropils- 
olfactory glomeruli, which receive and process olfac- 
tory signals from the antennae. 

Insect species with complex and flexible behavior 
possess well-developed mushroom bodies and anten- 
nal lobes, and larger insects have larger brains and 
more complex histological brain structure and gener- 
ally exhibit greater complexity of behavior (Goossen 
1949, Bernstein & Bernstein 1969). The largest mush- 
room bodies (relative to the rest of the brain) are 
found in social Hymenoptera. The morphological plas- 


194 


ant 


2S 


Z 
Ant.|bs. 


Fic. 1. Diagram of the butterfly brain showing the most important 
neuropils. Opt.lbs.—optic lobes, Ant.Ibs.—antennal lobes, mb— 
mushroom bodies, Ke—Kenyon cells, eb—central body, be—back of 
the eye, ant.n.—antennal nerve, ant.—antenna, es—esophagus. 


ticity of these brain structures has been demonstrated 
in bees (Withers et al. 1993, Winnington et al. 1996, 
Robinson 1998) and ants (Gronenberg et al. 1996). 
Mushroom bodies increase in size when these insects 
begin to perform complex and behaviorally more de- 
manding tasks. Neuropil growth related to behavioral 
changes has also been observed in non-social insects, 
such as fruit flies and rove beetles (Bieber & Fuldner 
1979, Technau 1984, Heisenberg et al. 1995). This 
growth was found to represent the further arborization 
and proliferation of existing brain cells, and not the 
production of new neurons. 

Flexibility of behavior and learning have been 
demonstrated in different species of Lepidoptera 
(Swihart & Swihart 1970, Papaj 1986, Weiss 1995, 
1997, Hartlieb 1996, Fan et al. 1997). Butterflies and 
moths have well-developed mushroom bodies (Ali 
1974, Sivinsky 1989), and large antennal lobes (Mat- 
sumoto & Hildebrand 1981). Both olfactory and visual 
learning have been described in Agraulis vanillae 
(Weiss 1995, Kroutov et al. 1999). 

Here we studied brain morphology in two groups of 
Agraulis. One group comprised butterflies collected in 
nature (“experienced” group) and the other group was 
reared and maintained in the laboratory in isolation 
from normal environmental stimuli (“naive” group). 
We investigated the hypotheses that the sizes of brain 
structures involved in information processing and 
learning vary according to the individual experience of 
butterflies, and that such structures should be larger in 
butterflies exposed to various environmental stimuli 
than in butterflies deprived of those. 


MATERIALS AND METHODS 


Adults and larvae of Agraulis vanillae were collected 
in Gainesville, Florida. All butterflies used in experi- 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


ments were collected during the 3-4 day period of the 
abundance peak of the species. Larvae were reared in 
the laboratory on their natural host-plant Passiflora in- 
carnata (L.), picked in the same area where the larvae 
were found. Laboratory reared adults spent 48 hours 
after eclosion in 25 x 25 x 25 cm screen cages. The lab- 
oratory conditions were 25°C, 65% relative humidity, 
L:D 16:8 h. Butterflies were fed a 25% sugar solution. 
For the preparation of the histological specimens 
butterfly heads were removed and fixed in Bouin's fix- 
ative, prepared 24 hours prior to usage, for 2 days. 
They were then rinsed in 70% ethanol and embedded 
in paraffin. Heads of 16 reared males, 10 reared fe- 
males, 17 wild males and 22 wild females were sec- 
tioned. The frontal microtome sections were 10 um 
thick and were stained with hematoxylin-eosin. 
Volumetric analysis was performed with an AIS/C 
image analysis system (Imaging Research, Inc.) inter- 
faced to a Zeiss Axiophot microscope via a Dage 72 
CCD camera. The following areas were measured in 
selected spaced sections on both sides of the brain: 
whole brain, antennal lobes, olfactory glomeruli, optic 
lobes, central body, mushroom body calyces, and the 
regions occupied by Kenyon cells. When areas were 
measured, this was done without awareness of the 
group to which that individual belonged. The volume 
of a brain structure was calculated using the formula 
Vol ovjecy = 2) A Xt XN 


(object) 

where n is the number of sections on which measure- 
ments were made, A is the area of a measured section, 
t is the distance between adjacent sections (e.g., sec- 
tion thickness), and N is the number of sections repre- 
sented by the section A,. Between 10 and 20 evenly 
spaced sections were used to determine the volume of 
each region. This corresponded to 50-100% of all the 
sections containing each measured structure. The rela- 
tive volume of each brain structure was calculated as a 
percentage of the volume of the whole brain. 

For statistical analysis of the data a fixed effects lin- 
ear model (ANOVA) was fit with PROC GLM (SAS 
v.8). That is, size was modeled as a function of the 
fixed effects ‘brain region’, ‘butterfly gender’ and ‘but- 
terfly group’ (“experienced”, “naive” and “control”). All 
relevant assumptions such as constant variance and 
normality were formally assessed. Due to the large 
number of multiple Bonferroni comparisons we tested 
at the 0.01 level of significance throughout. 

To exclude the possible effect of age on the changes 
in Agraulis brain, a control group of 10 males and 10 
females, reared in the laboratory was kept in cages for 
20-25 days after eclosion under the same conditions as 
described for the experimental group. The heads of 


VOLUME 56, NUMBER 4 


195 


a,b,c,d—tfrontal sections, e—sagittal section. Scale bars—100 um. 


control butterflies were sectioned, sections stained and 
brains measured as described above. 


RESULTS 


Figure 2 shows the sections of the measured brain 
structures in Agraulis vanillae. Most of the regions ex- 
hibit clearly defined boundaries. Because of the ab- 
sence of a clear boundary between the mushroom 
body's pedunculus and lobes, and the surrounding neu- 
ropil, attributable to the staining method chosen for the 
study, only mushroom body calyces were measured. 

Whole brain volume of Agraulis showed no signifi- 
cant variation according to group (Fig. 3). There was 
found to be a significant interaction of gender *group* 
brain region (p < 0.0001). Multiple pairwise compar- 
isons revealed the following patterns: “experienced” 
individuals of both sexes exhibited significantly larger 


mushroom bodies and olfactory glomeruli than did 
“naive” or “control” individuals (Fig. 4; Table 1). The 
relative volume of mushroom body calyces in “experi- 
enced” butterflies was greater than in “naive” ones by 
36% in males, and by 38% in females. Olfactory 
glomeruli were larger in “experienced” Agraulis by 
48% in males, and 24% in females. 

The Kenyon cells region and antennal lobes showed 
mixed outcomes. Within the Kenyon cells region, 
there were no significant differences in volume among 
the male groups, but “experienced” females exhibited 
smaller volumes than did “controls”. For the antennal 
lobes, “experienced” males have larger volumes than 
do “naive” males. There were no differences among 
the female groups. The central body and optic lobe re- 
gions exhibited no significant difference for any pair- 
wise comparison. 


196 


N 
1 


Volume (x1 0°um*) 


1 


EF NF CF EM NM cM 


Fic. 3. Whole brain volume in Agraulis vanillae. EF—“experi- 
enced” females, NF—‘“naive” females, CF—‘“control” females, 
EM—“experienced” males, NM—"naive” males, CM—“control” 
males. 


For most brain regions assessed, there was no sig- 
nificant difference between male and female volumes. 
However, the antennal lobes exhibited the following 
pattern: “naive” and “control” females have larger an- 
tennal lobes than their male counterparts (p < 0.0001 
in each case), but “experienced” females do not differ 
significantly from “experienced” males. 

For all brain regions, the “naive” and “control” 
groups exhibited no significant differences in volume, 
within males or females. 


DISCUSSION 


The results of our research demonstrate that in 
Agraulis differences in experience are correlated with 
changes in the volume of several of its brain regions in- 
volved in sensory information processing and memory 
formation. During their adult stage (2-4 weeks), 
Agraulis vanillae butterflies must perform various ac- 
tivities, the success of which can be enhanced by 
learning. Location of feeding sites with flowers that of- 
fer sufficient nectar reward, and recognition of poten- 
tial danger are of importance to both sexes. Female 
Agraulis need to find suitable host-plants on which to 
lay eggs. This involves not only recognition of the 
proper plant amongst a variety of other plants, but also 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


memory of the location of the host Passiflora patch, 
because butterflies of this species utilize vast habitats 
and linger at one spot for no longer than is necessary 
to complete either feeding or egg-laying. Males, in 
turn, need to locate the host-plant area to encounter 
females and mate. 

Detailed analysis of the captivity conditions and 
their specific influence on Agraulis’s experience, learn- 
ing and associated morphological changes in its brain 
was not attempted. However, it seems evident that 
captive laboratory-reared butterflies would have a 
greatly reduced range of external stimuli, being de- 
prived of space, visual stimuli, contacts with host- 
plant, flowers and sex partners. It was also impossible 
to determine precisely the age of “experienced” but- 
terflies, collected in nature. But because we collected 
them in a period of 10-14 days after the beginning of 
their abundance peak, we can estimate all collected 
butterflies to be of approximately the same age. 

Generally, measured brain structures were larger 
(relative to the volume of the whole brain) in “experi- 
enced” butterflies. But no difference in the relative 
volume of optic lobes and central body was recorded 
between two groups of Agraulis. The most dramatic 
increases in relative volume occurred in the mush- 
room bodies and olfactory glomeruli, whereas no size 
difference in optic lobes between “experienced” and 
“naive” butterflies was observed. Therefore, olfactory 
stimuli may be of primary importance in driving the 
structural changes in the Agraulis brain. Although but- 
terflies reputedly rely heavily on visual stimuli (Swihart 
1970, Silberglied 1979, 1984), and Agraulis is capable 
of visual as well as olfactory learning (Weiss 1995), 
their optic lobes only pass visual information to the 
central brain, where processing and integration of this 
information takes place. Therefore such a result is pre- 
dictable. 

The relative decrease in volume of the Kenyon cells 
region is rather hard to explain. However, because 
there was no change in the whole brain volume, this 
region’s relative decrease could represent an actual 
compression of the Kenyon cell clusters by the ex- 


TABLE 1. Relative volumes of brain regions as percentage of the whole brain volume in Agraulis vanillae. Within each box, different small 
case letters indicate significant differences, whereas the same letters indicate no difference. 


“Experienced” males 
“Naive” males 
“Control” males 


“Experienced” females 
“Naive” females 
“Control” females 


Mushroom 
body calyx 


Olfactory 
glomeruli 


2.01 + 0.08 a 
1.48 + 0.05 b 
1.38 + 0.07 b 


2.18 +0.10a 
1.58 + 0.05 b 
1.58 + 0.12 b 


1.45 + 0.09 a 
0.98 + 0.05 b 
0.98 + 0.09 b 


1.44 + 0.08 a 
1.16 + 0.05 b 
1.11 + 0.04b 


Kenyon cells Antennal Central Optic 
region lobes body Lobes 
0.69 + 0.07 a 3.74 + 0.35 b 0.61 + 0.09 a 64.4+3.0a 
0.75 + 0.05 a 3.41+0.10a 0.68 + 0.04 a 69.7+8.3a 
0.72 + 0.04 a 3.50 + 0.10 ab 0.66 + 0.05a 65.1+23a 
0.56 + 0.03 a 3.90 + 0.10 a 0.63 + 0.07 a 61.8+2.6a 
0.73 + 0.05 ab 4.00 + 0.20 a 0.64 + 0.03 a 62.4+42.2a 
0.84 + 0.08 b 400 +0.12a 0.66 + 0.02 a 6l.1+19a 


VOLUME 56, NUMBER 4 


Mushroom Body 


percent of whole brain 
percent of whole brain 


tl 


EM NM CM EF NF CF EM NM 


Antennal lobes 


Olfactory glomeruli 


CM EF NF CF 


Central body 


Si 


Kenyon cells 


0.4 4 


percent of whole brain 


EM NM CM EF NF _ CF 


Optic lobes 


percent of whole brain 
percent of whole brain 


EM NM CM EF NF _ CF EM NM CM 


percent of whole brain 


EF NF CF EM NM CM EF NF CF 


Fic. 4. Relative volumes of brain regions as percentage of the whole brain volume in Agraulis vanillae. EM—“experienced” males, NU— 
*naive” males, CM—“control” males. EF—“experienced” females, NF—‘naive” females, CF—“control” females. Different small case letters 
indicate significant differences, whereas the same letters indicate no difference for each sex. 


panding mushroom body calyces. This problem could 
be addressed by more detailed experimental analysis, 
for example assessment of cell packing density. 

The data presented here are similar in many ways to 
those of studies in other species of insects, which mea- 
sured the size differences in brain regions caused by 
different experience and behavioral repertoire. As in 
the present study, an increase in relative volume of 
mushroom bodies, and decrease in relative volume of 
the Kenyon cells region were reported for ants (Gro- 
nenberg et al. 1996) and bees (Withers et al. 1993). 
Also, there was no increase in the relative volume of 
the optic lobes in either of these insects. Olfactory 
glomerular volume was found to differ between 1-day- 
old and nurse bees (larger in nurses), but the increase 
was not maintained in foragers. For rove beetles 
mushroom body volume increase and no changes in 
optic lobe volume were recorded (Bieber & Fuldner 
1979). 

The sexual dimorphism found in the reorganization 
of some brain structures in Agraulis, namely the dif- 


ference in olfactory glomeruli volume between “naive” 
and “experienced” males being twice as great as that in 
females, can perhaps be explained by the differences 
in behavior of males and females. Females need to lo- 
cate host-plants and determine their suitability for 
oviposition, and males need to search for females and 
recognize proper chemical cues from suitable part- 
ners. Thus, each sex may rely on different environ- 
mental stimuli. The change in the intensity of these 
stimuli may effect butterflies of different sexes differ- 
ently, and cause the observed dissimilarity in the brain 
reconstruction. This dimorphism corresponds with our 
earlier findings in Agraulis learning (Kroutov et al. 
1999), where different learning capability was 
recorded for the two sexes. 

Measurements in the control group show that mor- 
phological changes in the brain of Agraulis vanillae are 
not age-related, but experience-related, since the rela- 
tive volumes of the studied brain structures in 2-day 
(“naive”) and 20-25-day (“control”) butterflies were 
not significantly different. These changes occur in only 


198 


a few brain compartments that are noted for their role 
in information processing and learning in insects. This 
further supports the hypothesis that growth of these 
brain regions is related to learning experience and be- 
havioral complexity of a butterfly. 

Further experiments involving the manipulation of 
various elements of the environment may lead to a 
better understanding of the exact relation between 
particular types of information, how they are pro- 
cessed, and changes they cause in the brain of Agraulis 
vanillae. It would be especially interesting to analyze 
the specific effects of various environmental “depriva- 
tions” and, inverted, the effect of additional stimuli on 
the changes in Agraulis’ brain structures. 


ACKNOWLEDGMENTS 


We thank Dr. M. S. Mayer for his helpful comments on our pa- 
per, Scott Whittaker for his help with the preparation of the illustra- 
tions and Galin Jones for helping us with the statistical analysis of 
our data. 


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Received for publication 6 March 2001; revised and accepted 21 
February 2002. 


Journal of the Lepidopterists’ Society 
56(4), 2002, 199-215 


PHYLOGENETIC ANALYSIS AND REVIEW OF PANACEA AND 
BATESIA BUTTERFLIES (NYMPHALIDAE) 


RYAN I. HILL 
Section of Integrative Biology, University of Texas, Austin, Texas 78712, USA 


Cara M. PENZ! AND P. J. DEVRIES 
Milwaukee Public Museum, 800 W Wells St., Milwaukee, Wisconsin 53233, USA 


ABSTRACT. Phylogenetic analysis of 53 morphological characters for five species of Panacea and Batesia hypochlora supports the sepa- 
ration of the two genera and showed that the monotypic genus Batesia is basal to Panacea. Male genitalia were uniform within Panacea and char- 
acters informative for phylogeny reconstruction were restricted to wing coloration. Illustrations of adults and genitalia, a brief diagnosis, and dis- 


tributions are provided for each species. 


Additional key words: _prola, procilla, regina, divalis, bleuzeni, chalcothea, lysimache, bella, hypochlora, Caryodendron, Euphorbiaceae. 


By possessing distasteful wings or body fluids, brightly 
colored butterflies are generally avoided by many verte- 
brate predators in nature. This phenomenon is particu- 
larly well known in various genera of Nymphalidae (e.g., 
Acraea, Heliconius, many Danainae and Ithomiinae), 
Papilionidae (e.g., Battus, Parides) and Pieridae (e.g., 
Mylothris, Delias, Appias, Perrhybris, Itaballia) among 
others (see Poulton 1908, Sywnnerton 1919, Carpenter 
1942, Fisher 1958, Chai 1986). Nevertheless, a great 
many of these same butterflies are eagerly sought after 
and prized by a different group of predators, human col- 
lectors. Although collector value may provide a metric of 
how garishly colored a particular butterfly might be, it is 
often a poor measure of how well we understand that 
species. Therefore, when considering biological or evo- 
lutionary understanding of particular butterflies, it is 
likely that drab ones are equally as well known as those 
that are brightly colored. Although well represented in 
museum collections, and available as virtual specimens 
on the internet, nymphalid butterflies in the genera 
Batesia Felder and Felder, 1862 and Panacea Godman 
and Salvin, 1883 are good examples of this phenomenon. 

The Neotropical genus Batesia occurs from central 
Colombia to eastern Ecuador, southeast Peru, western 
Brazil, and likely into northeast Bolivia; effectively an 
upper Amazonian distribution. On the other hand, 
members of Panacea are found from Costa Rica south 
across Venezuela and the Guianas, throughout the 
Amazon basin, and into Bolivia. 

Both Batesia and Panacea were originally described 
as monotypic genera, but only Batesia with its single 
species, hypochlora Felder and Felder, 1862 has re- 
mained so. The history of Panacea is somewhat convo- 
luted. Panacea prola (Doubleday, 1848) was initially 
designated the type species of Pandora Doubleday, 

' Adjunct professor at Pontificia Universidade Catélica do Rio 
Grande do Sul, Av. Ipiranga 6681, Porto Alegre, RS, 90619-900, 
Brazil. 


1848—a name used previously for different insect gen- 
era by at least seven different authors, and thus, an in- 
valid homonym (see Hemming 1967). In an attempt to 
settle this quandary, Kirby (1871) transferred all species 
of Pandora to Batesia. Godman and Salvin (1883), how- 
ever, felt that all species formerly in Pandora warranted 
separation from Batesia, and erected the genus Panacea 
to accommodate them—thus providing a panacea to the 
Pandora problem. Eight species have been described in 
Panacea—P. prola; P. procilla (Hewitson, 1852); P. 
regina (Bates, 1864); P. divalis (Bates, 1868); P. chal- 
cothea (Bates, 1868); P. lysimache Godman and Salvin, 
1883; P. bleuzeni Plantrou and Attal, 1986; and P. bella 
D’Abrera, 1987, not all that are currently regarded as 
valid species (see synonymies below). 

The vicissitudes of nomenclature aside, nearly all 
natural history studies suggest that Batesia and 
Panacea are distinct, but closely related genera. At 
present they are classified in the Biblidini along with 
Hamadryas, Ectima, Eunica, Myscelia, Dynamine, 
Colobura and other genera (Godman & Salvin 1883, 
Seitz 1916, Ackery 1984, Harvey 1991). 

Recent observations indicate that Batesia and 
Panacea share Caryodendron spp. (Euphorbiaceae) as 
host plants, and that their immature stages are very 
similar (DeVries et al. 1999). The correspondence of 
immature biology, classification, and the fact that these 
genera have never been assessed using cladistic meth- 
ods led us to ask whether B. hypochlora was separate 
from Panacea, or if it represented a derived species 
within Panacea. Accordingly, this study tests both hy- 
potheses through phylogenetic analysis of five species 
of Panacea plus Batesia hypochlora. Based on adult 
morphology we show that Batesia hypochlora is basal 
to Panacea, and that together they form a mono- 
phyletic group. We then present characters to aid in 
species identification, and provide notes relevant to fu- 
ture work on their taxonomy and natural history. 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 1. Batesia hypochlora, dorsal. Top row, males; bottom row, females. Left column, Garza Cocha, Ecuador; right column, Rondonia, Brazil. 


MATERIALS AND METHODS 


Species studied. Excepting P. chalcothea (see 
identification section below), our phylogenetic analysis 
included all valid species of Panacea (P. prola, P. 
procilla, P. regina, P. divalis, and P. bleuzeni) and Bate- 
sia hypochlora (Figs. 1-10). 

To assess intra-specific variation in wing pattern and 
genitalia, we examined specimens from five distinct lo- 
calities. Abundant material from a single site in eastern 
Ecuador (P. prola, n = 57; P. divalis, n = 55; P. regina, 
n = 43; and B. hypochlora, n = 24) allowed us to eval- 
uate morphological and phenotypic variation within a 
single population (see DeVries & Walla 2001 for site 
description). Whenever possible individuals from dif- 
ferent localities were dissected to evaluate morpholog- 
ical variation in the genitalia. Although a small number 
of specimens were available of P. procilla (n = 4) and P. 
bleuzeni (n = 2), these species are phenotypically dis- 
tinctive from other Panacea and characters could be 
scored with confidence. For P. bleuzeni, one specimen 


of each sex was used to score genitalia characters di- 
rectly, but wing and body characters were scored using 
the description of Plantrou and Attal (1986), the illus- 
trations in D’Abrera (1987:487, as P. bella) and pho- 
tographs from the private collection of G. Attal. Charac- 
ters 22 and 23 were scored as “missing” for P. bleuzeni 
due to lack of material. Table 1 lists the examined taxa, 
number of dissected individuals, and locality data. 

We used Biblis hyperia (Cramer, 1780) and 
Hamadryas arinome (Lucas, 1853), H. amphinome 
(Linnaeus, 1767), H. laodamia (Cramer, 1777), and H. 
feronia (Linnaeus, 1758) as outgroup taxa for phyloge- 
netic analysis. Based on larval and adult morphology, 
and host plant use (Euphorbiaceae) these taxa are con- 
sidered closely related to Batesia and Panacea (Seitz 
1916, Ackery 1984, Harvey 1991). 

Preparation of material. Genitalia were prepared 
with a standard treatment of 10% potassium hydroxide, 
examined with a stereomicroscope, and subsequently 
stored in glycerol. Illustrations are given in Figs. 11-13. 

Characters and terminology. Our character matrix 


VOLUME 56, NUMBER 4 


201 


Fic. 2. Batesia hypochlora, ventral. Left column, Garza Cocha, Ecuador; right column, Rondonia, Brazil. 


includes 53 characters (43 binary and 10 multistate), of 
which 24 were derived from males (23 from genitalia, 
one from wing coloration), 7 derived from females (6 
from genitalia and one from wing coloration), and 22 
from both sexes (16 from wing patterns, four from ve- 
nation, one from forelegs and one from body scales). 
Terminology for adult external morphology follows 
Scoble (1992). Terminology for male and female geni- 
talia follows Klots (1970) except for the use of hypan- 
drium and ramus, which follow the definitions in the 
glossary of Tuxen (1970) and Jenkins (1986, 1987, 
1990). We use hypandrium to mean “a male subgenital 
plate,” and ramus as “lateral or ventro-lateral process of 
male eighth sternite, directed posteriorly” (see glossary 
in Tuxen 1970; Jenkins 1983, 1986). In character 10 we 
follow D’Abrera (1987) where a “complete ocellus” 
consists of a spot surrounded by a round ring (e.g., P. 
procilla, Fig. 6), and an “incomplete ocellus” is a spot 
without a round outer ring (e.g., P. bleuzeni, Fig. 7). 
Phylogenetic analysis. We used a heuristic search 
in PAUP 3.1 (Swofford 1993) with all characters given 
equal weight, multi-state characters unordered, poly- 
morphic characters treated as exhibiting both states, 
and the search used a TBR branch swapping routine. 


Following analysis, Biblis hyperia was used to root the 
tree. Branch support was estimated by 500 bootstrap 
replicates, and we used MacClade 3.01 (Maddison & 
Maddison 1992) to identify character changes along 
the branches of the tree. The character list and data 
matrix are in Appendix | and 2. 


RESULTS 
Phylogeny 


Our analysis indicates that Panacea and Batesia are 
monophyletic, sister taxa. The single most parsimo- 
nious tree (tree length = 79, CI = 0.82, RI = 0.88) sug- 
gests that Batesia hypochlora is a sister species to 
Panacea, a relationship supported by four characters 
(Fig. 14; Table 2, clade 1). We found 11 autapomor- 
phies for B. hypochlora (Table 2, clade 2), and nine 
characters that justify the monophyly of Panacea 
(Table 2, clade 3). Our analysis also showed that all 
members of Panacea are morphologically similar, but 
they differ strongly from Batesia hypochlora. 

Among Panacea the genital morphology was notably 
conservative, and characters providing the basis for in- 
ferring species relationships were derived mostly from 


wing morphology. Only one male genital character (hy- 


202 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fic. 3. Panacea prola, dorsal and ventral. Top row, left, male; right, female. Bottom row, left male; right, female. All from Garza Cocha, Ecuador. 


pandrium, character 28) could be used to distinguish 
among Panacea species. However, as it represents an au- 
tapomorphy for P. divalis, character 28 was uninforma- 
tive for establishing phylogenetic relationships within 
Panacea. The grouping of P. regina, P. divalis, P. bleuzeni 
and P. procilla was supported by seven characters, all de- 
rived from wing pattern morphology (Table 2, clade 4). 
One character justified grouping P. divalis, P. bleuzeni 
and P, procilla (Table 2, clade 5) and a single character 
grouped P. bleuzeni and P. procilla (Table 2, clade 6). 


Identification and Taxonomy 


Here we provide synonymies, characters for identifi- 
cation of the study taxa, approximate geographical dis- 
tributions, and comments on phenotypic variation of 
the species included in our analysis. For completeness, 
we also provide taxonomic notes on P. chalcothea, al- 
though we did not examine this taxon directly. 


Batesia Felder and Felder, 1862 


Batesia Felder and Felder, 1862. Wien. ent. Monats. 
ILLS. 


Batesia hypochlora Felder and Felder, 1862 
(Figs. 1, 2, 1, 13) 


Batesia hypochlora Felder and Felder, 1862. Wien. 
ent. Monats. 6:113 

Batesia hypochlora hypoxantha Salvin and Godman, 
1868. Ann. Mag. Nat. Hist. (4)2:147 

Batesia hypochlora hemichrysa Salvin and Godman, 
1868. Ann. Mag. Nat. Hist. (4)2:147 

Batesia hypochlora chrysocantha Fruhstorfer, 1915. 
Soc. ent. 30(12):66 

Batesia hypochlora f. intermedia Michael, 1931. Ent. 
Zeit. 44(20):309-312 
Species characters. Forewing dorsal surface 

dark iridescent blue from basal to submedial areas, a 

prominent postmedial red band surrounded by 

black, apex iridescent blue. Hindwing dorsal surface 

mostly iridescent blue, with a postmedial black band 

and an iridescent blue marginal band from apex to 

tornus. Forewing ventral surface dark brown from 

basal to submedial areas and tornus, postmedial red 

band surrounded by brown, subapex yellow. Hind- 


VOLUME 56, NUMBER 4 


203 


Fic. 4. Panacea regina, dorsal. Top row, male; bottom row, female. All from Garza Cocha, Ecuador. 


wing ventral surface chalky yellow with a distinct 
black postmedial band and yellow marginal band 
from apex to tornus. 

Distribution. Western Amazonas, Brazil; Ecuador, 
Peru (Seitz 1916, D’Abrera 1987, Austin & Emmel 
1990, Robbins et al. 1996). 

Variation. Judging by the named subspecies (see 
synonomic list) the intensity of yellow on the ventral 
surface of the HW may vary. However, whether 
these names are biologically meaningful remains 
uncertain. We found little variation in our samples 
from Garza Cocha, Ecuador, although we note that 
Ecuadorian and Brazilian material differ in the 
respective width of the forewing subapical band 
(Big: 1): 


Panacea Godman and Salvin, 1883 


Pandora Doubleday, 1848. Gen. Diurnal Lep. p. 300 
Pl. 3 fig 5 

Panacea Godman and Salvin, 1883. Biol. Centr. Am. 
pp. 274-275 


Panacea prola (Doubleday, 1848) 
(Figs. 3, 11, 13) 


Pandora prola Doubleday, 1848. Gen. Diurnal Lep. p. 
300 Pl. 3 fig. 5 

Panacea prola female f. dubia Kretzschmar 1894. 
Deutsche ent. Zeit. “Iris” 6(2):158—160 

P. prola zaraja Fruhstorfer, 1912. Ent. Rundschau 
29(6):46 

P. prola amazonica Fruhstorfer, 1915. Soc. ent. 
30(12):66 

P. prola prolifica Fruhstorfer, 1915. Soc. ent. 30(12):66 

P. prola amazonica £. bronzina Bryk, 1953. Arkiv. Fur 
Zool. 5(1):1—268 
Species characters. Dorsal surface with broken 

blue-green iridescent bands. Forewing dorsal surface 

without a subapical line in both sexes, but some fe- 

males with a faint greenish-white subapical band. 

Hindwing dorsal surface without ocelli or blue sub- 

marginal line. Hindwing ventral surface bright red, 

generally without black markings, but sometimes with 

a faint black submarginal line. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fic.5. Panacea regina, ventral. Top row, male; bottom row female. All from Garza Cocha, Ecuador. 


Distribution. Panama, Colombia, Venezuela, 
Guianas and upper Amazon basin (Seitz 1916, 
D’Abrera 1987, Emmel & Austin 1990, Otero & 
Romero 1992, Lamas 1994, Robbins et al. 1996, Neild 
1996). 

Variation. We found wide variation in wing length, 
but little variation in color pattern in large samples 
from Garza Cocha, Ecuador. Small individuals appear 
to be the result of caterpillars feeding on poor quality 
Caryodendron leaves, or those that were semi-starved 
(pers. obs.). 

Subspecies. Panacea prola zaraja, from Venezuela, 
Merida; P. p. amazonica, from the upper Amazon; P. p. 
prolifica, from Ecuador. 


Panacea regina (Bates, 1864) 
(Figs. 4, 5, 11, 13) 
Pandora regina Bates, 1864. J. Entom. 2(10):213. 
Panacea regina victrix Fruhstorfer, 1915. Soc. ent. 
30(12):66. 
Species characters. Dorsal surface with broken 
blue-green iridescent bands. Forewing ventral surface 


with reddish apex and white subapical band but with- 
out the distinct red spots outlined by black in discal 
cell (see P. divalis). Hindwing dorsal surface with a 
blue medial band adorned with incomplete black 
ocelli that vary in size, and may reach the distal margin 
of the band; submarginal wavy line sometimes faint. 
Hindwing ventral surface red with broken submedial 
to medial transverse black lines, the most distal start- 
ing at Sc + Rs and ending at Cu,; faint post-medial 
ocelli in almost all cells; conspicuous black submar- 
ginal line. Females often with a short, white longitudi- 
nal stripe in ventral hindwing cell M,—M,, nearly at the 
center of wing. 

Distribution. Westem and upper Amazon 
(Ecuador, Peru, Brazil) (Seitz 1916, D’Abrera 1987, 
Lamas 1994, Robbins et al., 1996). 

Variation. In Ecuadorian and Brazilian samples we 
found that the medial ocelli on the dorsal hindwing vary 
considerably within populations. In females we found 
the ventral hindwing ocelli were sometimes incomplete. 

Subspecies. Panacea regina victrix, from Ecuador; 
see also P. chalcothea (below). 


VOLUME 56, NUMBER 4 


Jaret 


PPO UTUUCLT Lb (LiL coed Lab 
' “) ' © 


»! Sed 

Fic.6. Panacea chalcothea, male, dorsal and ventral, plus label. 
This specimen is an apparent syntype (see Identification and Taxon- 
omy). Note: whether chalcothea is a subspecies of P. regina or a valid 
species remains to be resolved. 


Panacea chalcothea (Bates, 1868) 
(Fig. 6) 


Pandora divalis Bates, 1868. Ent. mon. Mag. 4(44):170. 

This somewhat obscure taxon figures importantly in 
the history of Panacea, and its taxonomic status is un- 
resolved. Although we were unable to examine mate- 
rial of chalcothea directly, the photo provided by G. 
Lamas (Fig. 6) may serve as a starting point for identi- 
fying this taxon. Here we excerpt correspondence re- 
ceived from G. Lamas that bears directly on the taxo- 
nomic interpretation of Panacea chalcothea: 


“Bates (1868:170) described chalcothea based on at least 2 spec- 
imens, one female (?) illustrated by Hewitson ([1854], Ill. exot. Butts 
1: pl. [42], fig. 4), and thought by the latter to be the female of 
procilla; and one male from “southern Equador’. Hewitson’s “fe- 
male” belonged to the collection of the Entomological Society of 
London, and that specimen is almost certainly lost, while Bates’ 
male would have been in his collection, and should have gone to the 
BMNH through Godman and Salvin. There seems to be no Bates 
specimen of chalcothea from southern Ecuador at the BMNH. 
However, there is a male specimen from Bates’ collection, labeled 
chalcothea by Bates himself, but from “N Peru’, and I interpret this 
as a possible syntype of chalcothea, agreeing very well with the writ- 
ten description of the male given by Bates in his original paper. 


TaBLe 1. Number of dissected individuals and locality data. Ab- 
breviations for source collections are: P. J. DeVries (PJD); G. Austin 
(GTA); G. Attal (GA); Los Angeles County Museum (LACM); Mil- 
waukee Public Museum (MPM). 


Taxa Source of dissected material 


Ingroup 
Batesia hypochlora 2 males: Brazil (GTA) 

8 males: Ecuador, Sucumbios, Garza 
Cocha (PJD) 

1 female: Brazil (GTA) 

1 female: Ecuador, Sucumbios, Garza 
Cocha (PJD) 

1 male: French Guyana (GA) 

1 female: French Guyana (GA) 

5 males: Ecuador, Sucumbios, Garza 
Cocha (PJD) 

2 males: Brazil, Rondonia (GTA) 

3 females: Ecuador, Sucumbios, Garza 
Cocha (PJD) 

2 males: Brazil (n = 1) and Colombia 
(n = 1) (LACM) 

1 male: Colombia (MPM) 

1 female: Colombia (MPM) 

5 males: Ecuador, Sucumbios, Garza 


Panacea bleuzeni 


Panacea divalis 


Panacea procilla 


Panacea prola 


Cocha (PJD) 
3 females: Ecuador, Sucumbios, Garza 
Cocha (PJD) 
Panacea regina 5 males: Ecuador, Sucumbios, Garza 
Cocha (PJD) 
3 females: Ecuador, Sucumbios, Garza 
Cocha (PJD) 
Outgroups 
Biblis hyperia 1 male: Ecuador, Sucumbios, Garza 
Cocha (PJD) 
1 female: Ecuador, Sucumbios, Garza 
Cocha (PJD) 
Hamadryas amphinome 1 male: Ecuador, Sucumbios, Garza 
Cocha (PJD) 


1 female: Ecuador, Sucumbios, Garza 
Cocha (PJD) 

1 male: Ecuador, Sucumbios, Garza 
Cocha (PJD) 

1 female: Ecuador, Sucumbios, Garza 
Cocha (PJD) 

1 male: Ecuador, Sucumbios, Garza 
Cocha (PJD) 

1 female: Ecuador, Sucumbios, Garza 
Cocha (PJD) 

1 male: Ecuador, Sucumbios, Garza 
Cocha (PJD) 

1 female: Ecuador, Sucumbios, Garza 
Cocha (PJD) 


Hamadryas arinome 


Hamadryas feronia 


Hamadryas laodamia 


Bates may well have confused “S Ecuador” with “N Peru”. Anyway, 
that specimen from “N Peru’ most probably came from Amazonas 
department in Peru. . . . Now, [it] seems to me that chalcothea 
(based on Bates’ o.d. and the syntype referred to above) is . . . very 
probably a subspecies of regina, or could even be a full species. For 
the time being, I'm calling those 2 specimens as Panacea regina 
chalcothea, though I wouldn't be too surprised if they were to rep- 
resent a high altitude species distributed from Colombia to N Peru 
(if Hewitson’s “New Granada” locality for his specimen is correct, 


which is quite doubtful).” 


06 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 7. Panacea divalis, dorsal. Left column, males; right column, females. Top row, Rondonia, Brazil; middle and bottom rows, Garza 
Cocha, Ecuador. Note variation in medial bands and submarginal ocelli. 


Distribution. Apparently Western Amazonas Pandora divalis Bates, 1868. Ent. mon. Mag. 4(44):171. 


(Ecuador, Peru) and Colombia (?). Panacea procilla divalis Seitz, 1916. Die Gross 
Schmetterlinge der Erde p. 537. 
Panacea divalis (Bates, 1868) Species characters. Dorsal surface with broken 


(Higs: 7,8, 12513) iridescent blue-green bands. Forewing ventral sur- 


VOLUME 56, NUMBER 4 


207 


TABLE 2. Characters justifying the groupings of species and genera. MacClade 3.01 was used to map character changes on the most parsi- 
monious tree. Characters indicated in bold type were unique to the group they support (independent of reversals). 


Clade 1. Panacea and Batesia 


(2:0) Fringe of scales in forewing and hindwing outer margin solid dark color 
(16:0) Ventral surface of hindwing with black submarginal line that is discrete in anal area and more diffuse toward costal area 
(24:0) Thorax: ventral portion completely covered with red-orange scales 


(27:1) In lateral view: Hypandrium without anterior rod-like projections 


Clade 2. Batesia hypochlora 


(8:2) Males: Ventral surface of forewing apex dark, with a yellow band 


(19:0) Forewing venation: M, arched toward anal margin 


(25:0) Hypandrium: narrow, plate like, with obvious constriction near the middle of its long axis 


(29:0) In lateral view, anterior portion of tegumen extremely projected 


(30:1) Uncus tip in lateral view sharply hooked 
(32:1) Uncus short 


(33:0) In lateral/ dorso-lateral view, base of uncus with obvious large dorsal ridges 
(34:1) In lateral view, tip of uncus not reaching or extending beyond tip of valva 


(37:0) Distal portion of gnathos small and projected ventrally 


(43:1) Distal portion of valva with small bare chitinous tip 


) 

(38:0) In ventral view, distal portion of gnathos with a rounded invagination 
) 
) 


(53:0) Antrum mostly membranous 

Clade 3. Panacea 
(4:1) Forewing postmedial band expressed dorsally only 
(5:1) In dorsal view, forewing subapical white band reduced 
(7:0) Ventral surface of forewing with white subapical band 


(10:0) Ventral surface of hindwing largely colored red-orange, with or without purplish sheen 
(17:0) Ventral surface of hindwing with dark line imposed upon cross-vein m,—m, (at distal edge of discal cell) 


(23:0) Foreleg with white scales laterally 
(42:0) Distal portion of valva curving ventrally 


(44:0) In lateral view, basal portion of valva with large conspicuous ventrally produced rounded projection 
(46:1) In lateral view, distal portion of saccus straight to slightly projected upward 
Clade 4. Panacea procilla, Panacea bleuzeni, Panacea divalis and Panacea regina 
(8:0) Males: Ventral surface of forewing apex uniformly dirty red-orange 
11:0) Ventral surface of hindwing with prominent dark line across basal half of cell Sc + R, 
12:0) Ventral surface of hindwing with prominent dark line across discal cell 


14:0) Ventral surface of hindwing with nearly continuous line through medial area that crosses cells Sc + R,, Rs, M,, M,, M,, Cu, and Cu, 


( 
( 
(13:0) Ventral surface of hindwing discal cell with two black dots in basal half 
( 
( 


15:1) Ventral surface of Iniadlostag with dark line not contiguous and line in cell Cu, more apical than line in cell Cu, 
(18:0) Female: ventral surface of hindwing with white patch of scales in medial area of cell M, 


Clade 5. Panacea procilla, Panacea bleuzeni and Panacea divalis 


(5:0) In dorsal view, forewing subapical white band well developed 


(6:0) In ventral view, foaavoting discal cell with two red-orange spots, one at base and one at mid-length 


Clade 6. Panacea procilla and Panacea bleuzeni 


(3: 0) In dorsal view, male forewing with oblique, diffuse black band encroaching on postmedial blue/green band. 


face with reddish apex, white subapical band and dis- 
tinct red spots outlined by black in discal cell (see P. 
regina). Hindwing ventral surface brownish red with 
a faint purple sheen; broken transversal black medial 
lines, the most distal starting at Sc + Rs and ending at 
1A; postmedial ocelli (black “rings”) on almost all 
cells; conspicuous black submarginal line. Females 
with a short, white longitudinal stripe in ventral hind- 
wing cell M,—M,, nearly at the center of wing. In- 
complete Beclli ea dorsal surface of hindwing vary in 
size, and may be absent in some specimens. 

Distribution. Upper Amazon (Seitz 1916), Colom- 
bia to Peru (D’Abrera 1987) and western Brazil (Em- 
mel & Austin 1990). 

Variation. In males the dorsal hindwing marginal 


band varies among samples from Brazil and Ecuador; 

the dorsal hindwing ocelli vary from diffuse to sharp; a 

short, ventral longitudinal stripe may occur in ventral 

hindwing cell M,—M.. In females the white, ventral 

longitudinal stripe in Infcetoring cell M,—M, may be dif- 

fuse or faintly expanded into the two eal pou: 
Subspecies. None. 


Panacea procilla (Hewitson, 1852) 
(Figs. 9, 12, 13) 


Pandora procilla Hewitson, 1852. Exot. Butt. 1. 

Panacea lysimache Godman and Salvin 1883. Biol. 
Centr. Americana p. 275. 

P. procilla ocana Fruhstorfer, 1912. Ent. Rundschau 
29(6):46. 


OS JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 8. Panacea divalis, ventral. Left column, males; right column, females. Top row, Rondonia, Brazil; middle and bottom rows, Garza 
Cocha, Ecuador. Note variation in white stripe centered in cell M,—M,,. 


P. procilla salacia Fruhstorfer, 1915. Soc. Ent. 30(12):66. Species characters. Dorsal surface with broken 
P. procilla lysimache Seitz, 1916. Die Gross blue-green iridescent bands. Forewing ventral sur- 

Schmetterlinge der Erde p. 537. face with distinct red outlined by black in discal cell, 
P. procilla var. marmorensis Hall, 1917. Entomologist reddish apex and white subapical band. Hindwing 


50(651):171-174. ventral surface brownish red with a faint purple 


VOLUME 56, NUMBER 4 


209 


Fic. 9. Panacea procilla, dorsal and ventral. Left column, male; right column, female. Specimens from Cali, Colombia. 


sheen; broken transverse medial black lines, the most 
distal starting at Sc + Rs and ending at 1A; complete 
postmedial ocelli on almost all cells, those on cells 
M,—Cu, and Cu,—Cu, with iridescent pupil; conspic- 
uous black submarginal line. Dorsal surface of hind- 
wing with a medial blue band adorned with black 
ocelli; conspicuous submarginal wavy line. Females 
with white medial band on ventral forewing, and also 
with a white band on ventral hindwing from cell Sc + 
R,-Rs to M,—M,, sometimes interrupted on M,—M,. 

Distribution. Costa Rica south to Colombia and 
throughout the upper Amazon basin and the Guianas 
(Kretzschmar 1894, Apolinar 1926). 

Variation. We observed some males that have a 
short, white longitudinal stripe in ventral hindwing cell 
M,—M.,, nearly at the center of wing—a pattern similar 
to females of P. regina and P. divalis. 

Subspecies. Panacea procilla procilla, western 
Venezuela (Neild 1996), Pp. ocana, from lower Mag- 
dalena River, Colombia (Seitz 1916, D’Abrera 1987): 
P. p. salacia, from Colombia (Seitz 1916, D’Abrera 


1987); P. p. lysimache from Volcan Chiriqui, Panama, 
Finca la Selva, Costa Rica (DeVries 1987, 1989). 


Panacea bleuzeni Plantrou and Attal, 1986 
(Figs. 10, 12, 13) 


Panacea bleuzeni Plantrou and Attal, 1986. Bull. So- 
ciété Sciences Nat. 50:23. 

Panacea bella D’Abrera, 1987. Butterflies of the 
Neotropical Region, part II: p. 487, new syn- 
onoym 
Species characters. Dorsal surface distinctively 

blue or blue-green. Dorsal surface of hindwing with a 

blue medial band adorned with large black ocelli; wavy 

iridescent submarginal line conspicuous. Ventral 
forewing with distinct red outlined by black in discal 
cell, reddish apex and white subapical band (similar to 
procilla). Ventral hindwing with transverse medial 
black line continuous from cell Sc + Rs to vein 1A; 
ocelli faint. Females with white marking extending dis- 
tally along black medial line from cell Sc + Rs to 
Cu,-lA. 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 10. Panacea bleuzeni, female, dorsal and ventral. This figure is reproduced through the kind permission of B. d’Abrera [Butterflies of 
the Neotropical Region, part II1:487]. It is the type of Panacea bella D’Abrera, 1987. 


Distribution. Apparently endemic to the Guianas 
(Plantrou & Attal 1986). However, it’s overlapping 
range with procilla and close relationship to it (Table 
2, clade 6) suggest the possibility that this taxon may 
be a subspecies of procilla. This point needs critical 
evaluation. 

Synonymic notes. Examination of the collection 
of the BMNH by A. Neild (pers. com.) revealed that 
the single female holotype of P. bella is also a 
paratype of P. bleuzeni. This, therefore, indicates that 
P. bella and P. bleuzeni represent a single species with 
bella as a junior synonym of bleuzeni. Comparing the 
illustration of the type specimen of bella (in D’Abrera 
1987) with photographs of male and female P. 
bleuzeni provided by G. Attal confirms this assess- 
ment. 


DISCUSSION 


Our analysis showed that Batesia and Panacea form 
a monophyletic group, with B. hypochlora basal to 
Panacea. Therefore, despite similarities in early stage 
morphology and host plant use, we reject the hypoth- 
esis that B. hypochlora is a derived species from within 
Panacea. Our study confirms the maintenance of Bate- 
sia and Panacea as separate taxa (e.g., Godman & 
Salvin 1883, Seitz 1916), and serves as a framework for 
future systematic work on both genera. We note that, 
without examining material firsthand, P. chalcothea is 
presumed to be the sister taxon of P. regina. However, 
the phylogenetic position of chalcothea requires con- 
firmation, including its taxonomic rank. 

Insect genitalia are widely used for phylogenetic 


VOLUME 56, NUMBER 4 211 


procilla 


q 


bleuzeni 


divalis 


Fic. 11. Male genitalia: hypandrium, lateral view, ventral view (inset: tip of gnathos in ventral view). Panacea procila, P. bleuzeni, and P. divalis. 


reconstruction and delimiting species boundaries be- Panacea, however, we found that the genitalia were 
cause their morphology may diverge rapidly, and highly conserved and provided no informative char- 
therefore provide informative characters (Eberhard acters for phylogeny reconstruction, or discrimina- 


1985, Porter & Shapiro 1990, Arnqvist 1998). In tion among species. Rather, the species-level rela- 


regina 


prola 


hypochlora 


Fic. 12. Male genitalia: hypandrium, lateral view (inset: uncus in lateral view), ventral view (inset: tip of gnathos in ventral view). Panacea 
regina, P. prola, and Batesia hypochlora. 


to 
e 
bo 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


procilla divalis 


bleuzeni 


regina 


hypochlora 


Fic. 13. Female genitalia: ventral view, Panacea procila, P. divalis, P. regina, and P. prola. Lateral view: P. bleuzeni, and Batesia hypochlora 
(insets: genitalia in ventral view). Note differences in the number of ovarioles between P. bleuzeni and B. hypochlora. 


tionships proposed here were derived solely from 
characters of wing pattern (Fig. 14, Table 2). Our 
study suggests that the most distinctly colored 
species, P. prola, is basal to other congeners, with re- 
maining species groupings justified by differences in 
wing patterns. 

The distinctive behavior and coloration make 
Panacea easily recognizable in the field. However, in 
large samples from one Ecuadorian site we found con- 
siderable intraspecific variation in both genital mor- 
phology and wing color patterns. This concurs with 
Seitz (1916) who noted that in some Panacea species 
within population phenotypic variation may be greater 
than among population variation, indicating that there 
may be transitions among species with respect to color 
pattern. With the possible exception of P. prola, such 
phenotypic variation precludes the notion that sym- 
patric Panacea species can be positively identified in 
nature without capturing them. 

Batesia and Panacea are obvious and often abun- 
dant elements of many Neotropical butterfly faunas 


and museum collections. Nevertheless, some taxa are 
rare in collections, and this study points to several 
questions that will require a full taxonomic revision to 
resolve, particularly regarding the status of P. chla- 
cothea and P. bleuzeni. Although potentially useful 
tools for conservation ecology, little has been reported 
on the natural history Batesia and Panacea. What we 
do know is that adults of both genera show significant 
flight height preference in some lowland rainforests, 
and that trees in the genus Caryodendron are larval 
hostplants (see DeVries 1989, Montoya 1991, DeVries 
et al. 1999, DeVries & Walla 2001). We do not know if 
all taxa exhibit vertical stratification, if these butter- 
flies use other hostplant genera, or if some species are 
warningly colored (e.g., P. prola, Batesia) that repre- 
sent models in mimicry complexes. We believe that 
field studies, in concert with phylogenetic analyses of 
Hamadryas, Ectima, Eunica, and related genera is the 
next step toward understanding the evolution of Bate- 
sia and Panacea, and the diversification of the Bibli- 
dini. 


VOLUME 56, NUMBER 4 


Panacea procilla 
Panacea bleuzeni 
Panacea divalis 
Panacea regina 


Panacea prota 


Batesta hypochtora 


' Hamadryas laodamia 


Harmadryas arinome 


as 
Ihael 
mom Horpadryas feromia 


Hamadryas amphinome 


Pe 


comme PIDs Ay peria 


Fic. 14. Single most parsimonious tree obtained from the 
analysis of 53 characters for 11 species (tree length = 79, CI = 0.82, 
RI = 0.88). Numbers above and below tree branches represent boot- 
strap values and the number of unambiguous changes respectively. 


ACKNOWLEDGMENTS 


We thank Eric Schwartz and the staff of La Selva Jungle Lodge 
for supporting this and other aspects of our research. G. Attal 
(France), G. Austin (Nevada State Museum), and B. Brown (Natural 
History Museum of Los Angeles County) kindly facilitated our work 
by loaning specimens. We are sincerely grateful to P. Ackery (The 
British Natural History Museum), G. Lamas (Museo de Historia 
Natural, Universidad Nacional Mayor de San Marcos, Peru), G. Me- 
Gavin (Oxford University Natural History Museum), A. Neild (Lon- 
don), and C. Smith (The British Natural History Museum) for pro- 
viding historical information and comparing type specimens that 
helped unravel the taxonomic mysteries of P. chalcothea and P. 
bleuzeni. We thank B. d’Abrera for permission to reproduce his 
figures of Panacea bella. Comments by G. T. Austin and André 
Freitas improved the clarity of this manuscript. RIH thanks Lee 
Robertson for inspiration. This study was supported in part by the 
National Science Foundation (DEB 00-96241) and is dedicated to 
the memory of Sonny Clark, Bill Evans, Thelonius Sphere Monk, 
and Bud Powell. 


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MappIsoN, W. P. & D. R. Mappison. 1992. MacClade: version 
3.01. Sinauer, Sunderland. 

Montoya, D. C. 1991. Aspectos biologicos del gusano cachén del 
inchi (Panacea sp. possible prola) Revista Colomb, Ent. (Bo- 
gota) 17:41-45. 

NeILp, A. F. E. 1996. The butterflies of Venezuela Part I. 
Nymphalidae 1. Meridian Publications, Greenwich, London. 

OTERO, L. D. & F. RoMERO. 1992. Nota sobre la presences de 
Panacea prola Dbldy & Hew. (Lepidoptera: Nymphalidae) en el 
sector central de Cordillera de la costa Venezulana. Boletin de 
Entomologia Venezulana (N.S.) 7:171-172. 

PLANTROU, J. & S. ATTAL. 1986. Description Dune Nouvelle Es- 
pece Du Genre Panacea Decouverte En Guyane Frangaise: 
Panacea bleuzeni nov. sp. (Lepidoptera Nymphalidae). Bulletin 
de la Société Sciences Nat. 50:23-24. 

Porter, A. H. & A. M. Sapiro, 1990. The lock-and-key hypothe- 
sis: lack of mechanical isolation in a butterfly hybrid zone. En- 
tomol. Soc. Amer. 83:107—114. 

POULTON, E. B. 1908. Essays on evolution. Clarendon Press, Oxford. 


RosBINS, R. K., G. Lamas, O. H. H. MIELKE, D. J. Harvey & M. M. 
CASAGRANDE. 1996. Taxonomic composition and ecological 
structure of the species-rich butterfly community at Pakitza, 
Parque Nacional de Manu, Peru, pp. 217-252. In Wilson, D. E. 
& A. Sandoval (eds.), Manu, the biodiversity of southeastern 
Peru. Washington, D.C., Smithsonian. 

ScoBLE, M. J. 1992. The Lepidoptera: form, function and diversity. 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


SWYNNERTON, C. M. F. 1919. Experiments and observations bear- 
ing on the explanation of form and colouring, 1908-1913, 
Africa. J. Linn. Soc. (Zoology) 33:203-385. 

SWOFFORD, D. L. 1993. PAUP: phylogenetic analysis using parsi- 
mony, version 3.1.1. Sinauer, Sunderland. 

TuXEN, S. L. 1970. Taxonomist’s glossary of genitalia in insects. 
Munksgaard, Copenhagen. 


Seitz, A. 1916. 52. Genus Panacea. 


APPENDIX 1. 


British Museum (Natural History), London. 

53. Genus Batesia, pp. 
536-537. In Seitz, A. (ed.), Macrolepidoptera of the World. Vol. 
5. Alfred Kemen, Stuttgart. 


Received for publication 19 June 2002; revised and accepted 3 July 
2002. 


Character list used in the phylogenetic analysis. Relevant figures are noted, and comments are included when needed. Defini- 


tions are in the Characters and Terminology section. 


Wing Characters: 


aw we 


10. 


22. 


. Forewing postmedial band expressed dorsally and ventrally (0); expressed dorsally only (1 


. Ventral surface of forewing with white subapical band (0 
. Males, ventral surface of forewing apex: uniformly dirty red-orange (0) 
. Dorsal and ventral sides of hindwing consistently with four complete ocelli (0); dorsal side of hindwing with five noe. ocelli (lacking 


. Ventral surface of Inindbovinge discal cell with two black dots in basal half (0); devoid of such a pattern (1). Note: of the 57 
. Ventral surface of hindwing with: nearly continuous line through medial area that crosses cells Se + R,, Rs, M,, 


. Ventral surface of hindwing with: dark line in cell Cu, and cell Cu, contiguous (0); 


. Forewing outer margin: concave (0), straight (1), convex (2). 
. Fringe of scales in the outer margin of wings: solid dark color (0), dark interspersed with white sections (1). 
. In dorsal view, male forewing with oblique, diffuse black band encroaching on postmedial blue-green band (0 


); devoid of such a pattern (1). 
Note: P. bleuzeni was scored using original description, illustration in D’Abrera and photos provided by G. Attal. 

); absent or reduced (2). Note: H. laodamia and P. 
procilla were polymorphic for this character because of differences between the sexes. 


. In dorsal view, forewing subapical white band well developed (0); reduced (1); absent (2). 
. In ventral view, red-orange spots on forewing discal cell: 


two spots present, one at base and one at mid-length (0), one spot present, at mid- 
length (1), absent (2). 

); devoid of such pattern (1). 

; dark, same color as medial area (1); dark, with a yellow band (2). 


outer ring) and clearly separated from any black lines (1); ventral side of hindwing with four to six complete ocelli (2); devoid of such pat- 
terns (3). Note: To understand the variation in this character a large number of specimens were examined, and we ead no exceptions to 
the patterns described here (see Methods, Species studied). 

Ventral surface of hindwing largely colored red-orange, with or without purplish sheen (0); devoid of such a pattern (1). Note: although the 
presence of a purplish sheen has been used to separate P. procilla and P. divalis, we found this character to be present in both these species 
and variable within each of them. 


. Ventral surface of hindwing with prominent dark line across basal half of cell Se + R, (0); devoid of such a pattern (1). 
. Ventral surface of hindwing with prominent dark line across discal cell (0) 


); devoid a such a pattern (1). 

P. prola specimens ex- 
amined, three had two dots, 22 had one dot, and 32 lacked dots: in P. divalis, four of the 53 specimens had dots ee into a single marking. 
> Cu, mal Cu, (0); de- 
void of such a pattern (1). 

dark line not contiguous and line in cell Cu, more apical 
than line in cell Cu, (1); dark line not contiguous and line in cell C u, more basal than cell Cu, (2); ale k line absent from cell 1Cu, (3). 


. Ventral surface of hindlocng with black submar ginal line which is discrete in anal area and be comes more diffuse toward costal area (0); de- 


void of such a pattern (1). Note: P. bleuzeni was scored using the illustrations in D’Abrera (1987) and photos from the collection of G. Attal. 


. Ventral surface of hindwing with dark line imposed upon cross-vein m,—m, (at distal edge of discal cell) (0), devoid of such a dark line (1). 


Note: in P. prola, three of 53 specimens lacked the dark line. 


. Female, ventral surface of hindwing with white patch of scales in medial area of cell M, (0); devoid of white patch (1). Note: two males of P. 


procilla had similar white patch. In P. divalis one of 12 lacked the patch, and in P. regina two of 14 lacked the patch. 


. Forewing venation: M, arched toward anal mar gin (0); devoid of such a pattern (1). 
20. Forewing venation: M. arched toward anal margin (0); devoid of such a pattern (1). 
21. Forewing cross-vein m,—m, + cu,: joins M, + Cu, at or distal to the fork M, and Cu, ( 


0); proximally to the fork M, and Cu, (1); absent (2). 
Note: M, + Cu, denotes the combination a vein M, and Cu, proximal to tlie fork elie e they split. 


Forewing cross-vein r—m,, and the base of M, and ML inflated (0); not inflated (1). 


Body Characters: 


23: 
24. 


Foreleg with white scales laterally (0); devoid of white scales (1). 
Thorax: ventral portion completely covered with red-orange scales (0); devoid of such a pattern (1). 


Male Genitalia Characters: 


25. 
26. 
27. 


28. 


29 


Hypandrium: narrow, plate like, with obvious constriction near the middle of its long axis (0); broad, curling laterally, without a constriction (1). 
In lateral view, hypandrium with long ramus projecting posteriorly (0); devoid of projections (1). 
In lateral view, hypandrium with anterior rod-like projections (0); devoid of such a pattern (1). 


In lateral view, posterior corner of hypandrium extended into an obvious lobe-like process that projects dorsally (0); less lobe-like and not as 
projected dorsally (1). 


. In lateral view, anterior portion of tegumen extremely projected (0); devoid of such a pattern (1). 
30. 


In lateral view, uncus tip: pointed (0); sharply hooked (1). 


VOLUME 56, NUMBER 4 


31. Uncus: bifid (0); entire (1). 
32. Uncus: elongate (0); short (1). 


APPENDIX 1. Continued. 


33. In lateral/ dorso-lateral view, base of uncus with obvious large dorsal ridges (0); with small ridges (1); devoid of such a pattern (2). 
34. In lateral view, tip of uncus reaching or extending beyond tip of valva (0); devoid of such a pattern (1). 
35. Uncus with obvious, long setae dorsally (0); devoid of setae (1). 
36. Distal portion of gnathos: completely fused (0); bifid (1). 


37. Distal portion of gnathos: small and projected ventrally (0); large and projected posteriorly (1). 


38. In ventral view, distal portion of gnathos: with a rounded invagination (0); invaginated in a perfect “V” (1). 
39. Valva: with dentate process approximately 2/3 from its base (0); without such a process (1). 
40. Process of valva: projecting dorsally (0); projecting medially (1). 
4]. Process of valva: with setae (0): without setae (1). 
42. Distal portion of valva: curving ventrally (0); curving dorsally or straight (1). 


43. Distal portion of valva with large bare chitinous tip (0); with small bare chitinous tip (1); devoid of such pattems (2). 
44. In lateral view, basal portion of valva: with large conspicuous ventrally produced rounded projection (0); devoid of such a pattern (1). 


45. In lateral view, rod-like projections of juxta: large (0); small (1). 
46. In lateral view, distal portion of saccus: strongly projected upward (0); straight to slightly projected upward (1). 
47. In lateral view, vinculum with obvious dentate process along anterior margin (0); process shaped as a bump, not dentate (1). 


Female Genitalia Characters: 
48. Signa: present (()); absent (1). 
49. Sterigma: present (0); absent (1). 


50. Lamella antevaginalis: continuous across ventral surface (0); split (1). 

51. Lamella antevaginalis: fused to edge of eighth sternite (0); not fused (1). 
52. Ductus seminalis connecting to ductus bursa: very near corpus bursa (0); far from corpus bursa, and near ostium bursa (1). 
53. Antrum: heavily sclerotized (0); mostly membranous (1). 


Ingroup 
Batesia hypochlora 
Panacea prola 
Panacea procilla 
Panacea divalis 
Panacea regina 
Panacea bleuzeni 

Outgroups 
Biblis hyperia 
Hamadryas laodamia 
Hamadryas arinome 
Hamadryas amphinome 
Hamadryas feronia 


APPENDIX 2. Character Matrix. 
1010221231 1111301101 011001101 
0011100130 1111300111 0100111010 
000(0,1)000000 0000100011 0100111010 
0011000030 0000100011 0100111110 
0011120020 0000100011 0100111010 
0002000010 0010000011 ??00111010 
2112221131 1111311111 2111110?10 
211(0,2)221131 JI11311111 1011100?10 
1110221131 1111311110 1001100?10 
1110271130 1111310110 1001100?10 
111?211?01 0110210710 1001100710 


1101110000 
101011111? 
101011111? 
101011111? 
1010111112 
101011111? 


0020101?1? 
1010011201 
1011011101 
10?1011101 
1011011101 


1111001111 
?000011110 
?000011110 
?000011110 
?000011110 
?000011110 


?101001100 
0121100011 
0121100011 
0121100011 
0121110011 


215 


iL 
110 
110 
110 
110 
110 


110 
000 
000 
000 
000, 


Journal of the Lepidopterists’ Society 
56(4), 2002, 216-233 


REVISED IDENTITIES AND NEW SPECIES OF AETHES FROM 
MIDWESTERN NORTH AMERICA (TORTRICIDAE) 


MICHAEL SABOURIN 
23476 Johnson Rd., Grantsburg, Wisconsin 54840, USA, mothvet@grantsburgtelcom.net 


WILLIAM E.. MILLER 
Department of Entomology, University of Minnesota, 1980 Folwell Ave., St. Paul, Minnesota 55108, USA, mille014@tc.umn.edu 


Eric H. METZLER 
1241 Kildale Square North, Columbus, Ohio 43229, USA, spruance@infinet.com 


JAMEs T. VARGO 
13890 Ireland Road, Mishawaka, Indiana 46544, USA, Jvargo51@aol.com 


ABSTRACT. Ongoing systematic study of Nearctic Tortricidae revealed several historical misidentifications and new species of Aethes in 
the Midwest. We redefine A. promptana (Robinson), A. angulatana (Robinson), A. argentilimitana (Robinson), and A. atomosana (Busck); res- 
urrect A. interruptofasciata (Robinson); and synonymize A. labeculana (Robinson) with the prior A. argentilimitana, and A. sublepidana (Kear- 
fott) with the prior A. interruptofasciata. We describe as new species A. sexdentata Sabourin & Miller, A. razowskii Sabourin & Miller, A. 
westratei Sabourin & Miller, A. matheri Sabourin & Miller, A. terriae Sabourin & Miller, A. baloghi Sabourin & Metzler, and A. matthew- 
cruzi Sabourin & Vargo. The status of six species previously considered incertae sedis is resolved, and the number of recognized Nearctic Aethes 
species is increased from 27 to 34. Conclusions are based on type study as well as on more than 500 pinned specimens and more than 300 gen- 
italia preparations in 29 museums and private collections. A comprehensive definition of Aethes also is presented. 


Additional key words: Tortricinae, Cochylini, taxonomy. 


The genus Aethes is holarctic, with 27 species previ- 
ously recognized in North America (Metzler 1999, 
Pogue 1986, Powell 1983, Razowski 1986, 1994, 1997). 
The higher category to which Aethes belongs has been 
ranked variously from tribe to family, but a concensus 
is emerging that the appropriate rank is tribe, namely 
Cochylini of Tortricinae. 

Cochylini are difficult to discriminate and identify 
because of sibling or cryptic species, variability in mac- 
ulation and size, and geographic variation. Lack of de- 
tailed systematic work, including unstudied types and 
mixed type series, adds to the confusion surrounding 
the taxonomy of Aethes and its relatives. 

In the early 1900's, Busck (1907) and Kearfott 
(1907a, 1907b) described a number of Cochylini 
species. Forbes (1923) provided a synopsis of species 
recorded from New York and neighboring areas. With 
additional collecting and increased interest in biodiver- 
sity, Cochylini have attracted new attention. In a dis- 
sertation, Pogue (1986) proposed a new generic classi- 
fication of Nearctic Cochylini, including Aethes, and we 
draw liberally upon his work, all of which has not been 
published. He recognized 102 described species in 23 
genera, with 8 species incertae sedis. Razowski (1997) 
recently reviewed Canadian species, but did not exam- 
ine all types. Metzler (1999) described two species in 
connection with work on Midwestern prairie insects. 


Structurally, Aethes has one notable synapomorphy: 
the paired sicklelike structures of the male socii. Oth- 
erwise, Aethes includes markedly diverse genital struc- 
ture. A number of structural characters useful for 
species discrimination are found on or in the aedeagus 
and associated parts. 

This paper redefines and illustrates five Aethes 
species whose identities have been confused histori- 
cally. In addition, seven new species are added to the 
Nearctic fauna. The seven new species raise the 
number of Aethes recognized in North America from 
27 to 34. 


MATERIALS AND METHODS 


This study is based on more than 500 pinned 
adults assembled from 29 sources listed at the end of 
this section. Genitalia of more than 300 specimens 
were prepared. For the five previously known 
species treated here, “Specimens examined” sections 
are shortened to States and Provinces of specimen 
origin, months of capture, and depositories. Detailed 
lists of such specimens are available from the au- 
thors. 

Wing lengths were measured by caliper under mag- 
nification to the nearest 0.2 mm. Colors were deter- 
mined under incandescent light by comparison with 
the swatches of Smithe (1975). 


VOLUME 56, NUMBER 4 


i) 
a 
+l 


OM fascia 


TM facia Ml 


Spined harpe 


1b 


Lamella 
postvaginalis 


il 


Ventral margin 2b 


Posterior 
margin 


Ne” -2c 38 


Ic Id * 2c 2d 

Fics. la-3d. Aethes species. la. Wings of A. promptana lectotype ¢. 1b, Genitalia of A. promptana lectotype ¢ (prep. MGP936). 1c, Aedea- 
gus of A. promptana lectotype ¢ (prep. MGP936). 1d, Genitalia of A. promptana 2 from Grantsburg, Wisconsin (prep. MS99182). 2a. Wings of 
A. angulatana d from St. Joseph Co., Indiana. 2b, Genitalia of A. angulatana 6 from St. Joseph Co., Indiana (prep. MS00253). 2c, Aedeagus of 
A. angulatana ¢ from St. Joseph Co., Indiana (prep. MS00253). 2d, Genitalia of A. angulatana lectotype ° (prep. MGP932). 3a. Wings of A. ar- 
gentilimitana 6 (A. labeculana lectotype 2). 3b, Genitalia of A. argentilimitana 6 (A. labeculana lectotype 3) (prep. MGP934). 3c, Aedeagus of 


A. argentilimitana lectotype ¢ (prep. MGP931). 3d, Genitalia of A. argentilimitana 2 from Grand Isle, Vermont (prep. MS01052). 


Genitalia were prepared by placing abdomens in 
cold 10% KOH for 24-48 hours. After removal from 
KOH, they were cleaned in distilled water, soaked in 
70% isopropyl alcohol, then stained in aqueous Chlo- 
razole Black E (3 minutes for males and 1—2 minutes 
for females). Following Chlorazole staining, pelts were 
double-stained with lignin pink and acid fuchsin 
(males for 3 minutes, females for 1 minute). Pelts were 
then returned to 70% isopropyl alcohol, and the geni- 
talia removed. The male aedeagus was separated with 
the juxta remaining attached to it. Genitalia then were 
placed in 99% isopropyl alcohol for 15 minutes. Fi- 


nally, they were stored in glycerine in glass pin vials, or 
mounted on slides in Canada balsam or Euparal 
thinned with Cellosolve, and dried at 19°C for five 
weeks. Vinyl cover-slip props were used on female 
slides to reduce distortion. 

Whole specimens were photographed under tung- 
sten lights with a 35-mm camera attached to a dissect- 
ing microscope, except for Fig. 1la which was pro- 
duced with a Polaroid microscope camera with a fiber 
optic light source. Genitalia were photographed with a 
35-mm Olympus photomicrographic apparatus. All 
photo images were scanned, after which they were en- 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Cervix 
bursae 


sd 7 & 


Fics. 4a-6d. Aethes species. 4a. Wings of A. interruptofasciata 3 from Burnett Co., Wisconsin. 4b, Genitalia of A. interruptofasciata 3 from Lk. 
Katherine, Wisconsin (prep. MS01040). 4c, Aedeagus of A. interruptofasciata ¢ from Allegheny Co., Pennsylvania (prep. MSO1056). ). 4d, Genitalia of 
A. interruptofasciata ? from Kanawha Co., West Virginia (prep. MS99068). 5a, Wings of A, sexdentata paratype ¢ from S. Burlington, Vermont. 5b, 
Genitalia of A. sexdentata holotype ¢ from Grand isle, Vermont (prep. MS10046). 5c, Aedeagus of A. sexdentata holotype 6 oan Grand Isle, Ver- 
mont (prep. MS10046). 5d, Genitalia of A. sexdentata paratype ° from Clark Co., Illinois (prep. MS98427). 6a, Wings of A. razowskii paratype ¢ from 
Allegan Co., Michigan. 6b, Genitalia of A. razowskii holotype ¢ from Schoolcraft Co., Michigan (prep. MS01050). 6c, Aedeagus of A. raz owskii 
paratype ¢ from Wineton Co., Alabama (prep. MS99196). 6d, Genitalia of A. razowskii par atype 2 from Winston Co., Alabama (prep. MS99197). 


hanced and cropped with Apple Photoflash software, 
then laser printed. 

Abbreviations used are as follows: AL, at light; BL, 
black light; BLT, black-light trap; FW, forewing; gen. 
prep., genitalia preparation; HW, hindwing; IG, in 
glycerine; IM fascia, inner median fascia (antemedian, 
antemedial, and median of authors); LT, light trap; 
OM fascia, outer median fascia (postmedian or preapi- 
cal of authors); n, number of specimens on which a 
statement is based; ND, no date; ST line, subterminal 
line; UV, UVL, ultraviolet light. Collection dates are in 
month/day/year format. 


Certain anatomical terms are defined or illustrated or 
both as follows. Anal margin: the straight portion of the 
forewing trailing edge (Fig. 2a); anal crescent: mark on 
anal margin of the forewing (Fig. 2a); harpe: slightly 
raised spined area on the interior distal half of the valva 
(Fig. 1b); costa of the valva: dorsal margin of the valva 
(Fig. 2b); posterior and ventral margins of the valva (Fig. 
2b); phallobase: sheath surrounding the base of the 
aedeagus (Fig. 4c). Illustrations of some other terms are 
noted on first use in the text. 

Collection abbreviations used are: AMNH, Ameri- 
can Museum of Natural History, New York, New York; 


VOLUME 56, NUMBER 4 


Te 7d 


219 


Female 
unknown 


Fics. 7a-9c. Aethes species. 7a, Wings of A. westratei paratype ° from Cass Co., Michigan. 7b, Genitalia of A. westratei paratype d from 
Cass. Co., Michigan (prep. MS01048). 7c, Aedeagus of A. westratei paratype ¢ from Cass. Co., Michigan (prep. MS01039). 7d, Genitalia of A. 
westratei paratype 9 2 from Cass Co., Michigan. (prep. MS98146). 8a, Wings of A. matheri holotype 4 om Long Beach, Mississippi. 8b, Geni- 
talia of A. matheri paratype ¢ from Hinds Ca, MsSissigya (prep. MSO1082). 8c, Aedeagus of A. matheri paratype d 5 from Hinds Co., Mississippi 
(prep. MS01082). 8d, Genitalia of A. matheri paratype ° from Harrison Co., Mississippi (prep. MS01132). 9a, Wings of A. terriae holotype 4 


from Kalamazoo Co., Michigan. 9b, Genitalia of A. terriae paratype 6 


paratype 3 from Cass Co., Michigan (prep. MS01088). 


ANSP, Academy of Natural Sciences of Philadelphia, 
Philadelphia, Pennsylvania; BGS, B. G. Scholtens col- 
lection, Charleston, South Carolina; BM, The Natural 
History Museum, London, England; BMMS, B. 
Mather collection, Mississippi Entomological Mu- 
seum, Mississippi State, Mississippi; CMNH, Carnegie 
Museum of Natural History, Pittsburgh, Pennsylvania; 
CNC, Canadian National Collection of Insects, Ot- 
tawa, Ontario; EME, Essig Museum of Entomology, 
University of California, Berkeley; FMNH, Field Mu- 


S from Cass Co., Michigan (prep. MS01088). 9c, Aedeagus of A. terriae 


seum of Natural History, Chicago, Illinois; GJB, G. J. 
Balogh collection, Portage, Michigan; FMPS, Frost 
Entomological Museum, Pennsylvania State Univer- 
sity, University Park; INHS, Illinois Natural History 
Survey, Champaign; JDG, J. D. Glaser collection, Bal- 
timore, Maryland; JHW, J. H. Wilterding III collec- 
tion, East Lansing, Michigan; JRH, J. R. Heitzman 
collection, Independence, Missouri; JV, J. Vargo col- 
lection, Mishawaka, Indiana; MAR, M. A. Roberts col- 
lection, Steuben, Maine; MS, M. Sabourin collection, 


Anellus 
lobe 


0c 10d 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fics. 10a-12d. 10a, Wings of A. baloghi holotype ° from Yonkers, New York. 10b, Genitalia of A. baloghi paratype ¢ from Allegan Co., 
Michigan (prep. MS01083). 10c, Aedeagus of A. baloghi paratype ¢ from Newaygo Co., Michigan (prep. MS01098). 10d, Genitalia of A. baloghi 
paratype ° from Whitesbog, New Jersey (prep. MS01077). 1la, Wings of A. atomosana ¢ from Kings Co., Nova Scotia. 11b, Genitalia of A. ato- 
mosana 3 from Lunenburg Co., Nova Scotia (prep. MS01047). llc, Aedeagus of A. atomosana ¢ from Lunenburg Co., Nova Scotia (prep. 
MS01047). 11d, Genitalia of A. atomosana ° from Bumett Co., Wisconsin (prep. MS00653). 12a, Wings of A. matthewcruzi holotype ¢ from 
Roseau Co., Minnesota. 12b, Genitalia of A. matthewcruzi holotype ¢ from Roseau Co., Minnesota (prep. MS00167). 12c, Aedeagus of A. 
matthewcruzi holotype ¢ from Roseau Co., Minnesota (prep. MS00167). 12d, Genitalia of A. matthewcruzi paratype ° from Burnett Co., Wis- 


consin (prep. MS00342). 


Grantsburg Wisconsin; MSG, M. S. Griggs collection, 
Grand Isle, Vermont; MSUC, Michigan State Univer- 
sity, East Lansing; NSPM, Nova Scotia Museum, Hal- 
ifax; RL, R. Letsinger collection, Sarcoxie, Missouri; 
UMMZ, University of Michigan, Ann Arbor; UMRM, 
University of Missouri, Columbia; UMSP, University 
of Minnesota Entomology Museum, St. Paul; USNM, 
National Museum of Natural History, Washington, D. 
C.; UWEM, University of Wisconsin, Madison; WPW, 
W. P. Westrate collection, Cassopolis, Michigan; YPM, 


Yale University Peabody Museum of Natural History, 
New Haven, Connecticut; ZMUH, Zoological Mu- 
seum, University of Helsinki, Finland. 


Genus Aethes Billberg, 1820 


Type species: Pyralis smeathmanniana Fabricius, 
1781, designated by Fernald (1908). 
The characterization that follows is based mainly on 
North American species. 
No significant external sexual dimorphism. Speci- 


VOLUME 56, NUMBER 4 


mens cannot reliably be sexed by number of frenular 
bristles; males have one bristle, females one or two. 
Head. Proboscis conspicuous; ocelli present; scales 
of labial palpus expanded ventrally beyond third seg- 
ment, inner side usually of FW ground color, outer 
side of FW marking colors. Thorax. Short dorsal tuft 
usually present; lateral tufts of metanotum consisting 
of flattened hairs or elongate scales. FW longer than 
wide, lacking costal fold, costa slightly curved, termen 
slanted toward body, apex acute or obtuse. All veins 
usually free, R, and Cu, originating, respectively, from 
middle and distal 4 of discal cell; origins of R,, R,, 
R,, M,, and M, usually closer to veins following them 
than to veins preceding them, R, extending to costa 
before apex, origins of R, and M, usually closer to 
veins preceding them than to those following them, 
M, and Cu, adjacent or connate, CuP absent, the last 
probably constituting a cochyline synapomorphy 
(Pogue 1986); a vestigial chorda present in some 
species. Variably developed subcostal and cubital reti- 
nacula retain female frenular bristles. Ground color 
usually pale with markings darker; basal fascia usually 
thin, sometimes broad, or absent; median fascia re- 
duced, represented by inner and outer parts, the in- 
ner one sometimes complete from costa to anal mar- 
gin (Fig. 2a), sometimes divided into costal and anal 
parts, the anal part forming a crescent along the anal 
margin; outer fascia complete in some species, in oth- 
ers represented by a small spot along costa; discal spot 
usually present, consisting of scales at the distal end of 
the discal cell beyond the inner median fascia; ST line 
(Fig. 2a) occasionally present. HW costal fold usually 
lacking, veins Rs and M, stalked; origins of M, and 
Cu, well separated. Abdomen. Proximal apodeme of 
sternum 2 a short stub or cone. Anterolateral process 
longer than proximal apodeme. Venulae of sternum 2 
long, contrasting. Male genitalia: Aedeagus variable 
in length and width, vesica may have one large commu- 
tus accompanied by a small bundle of cornuti, or cor- 
nuti may be absent altogether. Anellus lobes (Figs. 
10c, lle, 12c) sometimes present. Juxta (Figs. 8c, 12c) 
typically a simple subquadrate or subrectangular plate 
attached to apex of phallobase, caulis absent. Valva 
broad, posterior margin (Fig 2b) variable, some 
species with dentiform spines on the harpe (Fig. 1b) 
in a pattern that makes the distal portion of the valva 
resemble a shark snout. Sacculus varying in length 
and shape. Uncus usually indiscernible, socii with 
dual apical sicklelike structures. Apex of vinculum 
arms variable in size, free, or fused by a thin mem- 
brane. Female genitalia: Papillae anales elongate, 
narrow, lightly sclerotized. Anterior and posterior 
apophyses slender, as long as or longer than papillae 


anales. Sterigma variable, sometimes reduced; with- 
out bilateral sclerotized bands connecting sterigma to 
anterior apophyses; lamella postvaginalis (Fig. 1d) 
usually present as a medial process or plate. Antrum 
robust to reduced. Ductus bursae highly variable, 
ranging from absent to present to coalesced with cor- 
pus bursae such as to be almost unrecognizable. 
Cervix bursae (Fig. 5d) sometimes present. Supple- 
mentary bursa (accessory bursa of authors) present, 
usually originating dorsally at base of antrum. Ductus 
seminalis originating from ductus-, cervix-, or corpus 
bursae, usually not from supplementary bursa. 
Signum (Fig. 10d) usually absent, but if present rep- 
resented by sclerotized vestiges of ductus bursae or 
broad sclerotized plates. 

Yasuda (1972) mentions peculiar bristles on tarsal 
segments 1-4 of the hind leg as being of taxonomic im- 
portance. These are short, dark spines occurring on 
the ventral surface of tarsomeres, usually singly or in 
rows at joints. In Aethes species treated here, the bris- 
tle groups on apical edges of eutarsal and distitarsal 
segments occur in a 3:3:3:3 pattern. 


Aethes promptana (Robinson) 


(Fig. la-d) 


Conchylis promptana Robinson (1869:286, pl. 8, fig. 
80) (Lectotype 6 designated here, type # 7440, #77, 
no data, FW 6.0 mm long, gen. prep. MGP936 [ Fig. 
1b, c], in ANSP). Fernald (1882:25), Handfield et al. 
(1997:44). 

Phalonia promptana; Dyar et al. (1903:487), Forbes 
(1923:508, in part), McDunnough (1939:60), 
Brower (1983:50, in part). 

Phtheochroa promptana; Poole & Gentili (1996:876). 

Aethes promptana; Razowski (1997:124, figs. 150-152, 
in part). 

Male and female exterior. Frons cream colored, 
vertex, notum, and tegulum buff-yellow; labial palpus 
3x longer than horizontal eye diameter. Thoracic tuft 
inconspicuous. Hind leg buff except for the shorter 
tibial spurs which are darker. FW (Fig. la) 5.7-7.9 
mm long (n = 21). Ground color cream, markings 
buff-yellow with scattered brown scales; fasciae not 
bordered by darker scales; costa and anal margin out- 
lined with short striae; basal fascia extending to costa, 
also slightly into discal cell at midpoint; IM fascia (Fig. 
2a) extending into cell, coalescing with discal spot; anal 
crescent and OM fascia (Fig. 2a) present; ST line (Fig. 
2a) extending to apex; fringe of ground color. HW drab 
gray. 

Male genitalia (Fig. 1b, c) (n = 7). Costa of valva 
straight, posterior margin broad, rounded, numerous 
dentiform spines on harpe; sacculus short, ventral 


margin straight. Median process of transtilla broad, tri- 
angulate, with a toothlike point. Aedeagus thick, vesica 
bearing one large, thick cornutus, a bundle of smaller 
cornuti, and a distal crescent-shaped sclerite. Anellus 
lobes absent. 

Female genitalia (Fig. 1d) (n = 9). Lamella post- 
vaginalis a large triangular medial process; anterior 
apophyses longer than posterior; antrum robust, elon- 
gate, tapering on anterior half. Cervix bursae sinistro- 
caudad of corpus bursae when viewed as in Fig. Id. 

Diagnosis. Superficially resembling A. patricia 
Metzler, but the FW markings of A. promptana are 
buff rather than red. Also, the posterior margin of the 
valva, the sacculus, median process of transtilla, and 
aedeagus (Fig. 1b, c) are all broader than in A. patricia 
(Metzler 2000: fig. 2a, b); the valval posterior margin is 
simple rather than edged with dense setae as in A. pa- 
tricia, and the female stemum 7 (Fig. 1d) is not heav- 
ily sclerotized as in A. patricia (Metzler 2000: fig. 2c). 

Discussion. Razowski’s (1997) concept of A. 
promptana apparently is based on a misidentified 
specimen. He did not report examining the specimen 
designated here as lectotype, which is undoubtedly 
the Pennsylvania male referred to by Robinson 
(1869), and thus a syntype. Although Klots (1942) 
never designated a lectotype, he mentions selecting 
Robinson’s “best specimens” for the purpose. We des- 
ignate a lectotype to prevent confusion in the future 
about the identity of this taxon. Razowski’s A. promp- 
tana is described later on here as A. razowskii, new 
species. 

Biology. Larval foodplants unknown. April—July 
adult captures suggest a univoltine or bivoltine life 
cycle. 

Specimens examined (n = 21, including lectotype 

3). Counties and months of capture by State and 
Province: Canada. NOVA SCOTIA: Cumberland, July; 
Queens, June (CNC, NSRM, UMSP). USA. ILLINOIS: 
Putnam, May (INHS). MAINE: Washington, July 
(MAR). MISSISSIPPI: Oktibbeha, April; Tishortaw, 
May; Warren, April (UMSP. BMMS). MISSOURI: 
Boone, May (UMRM). NEW JERSEY: Arlington, Au- 
gust (ANSP). OHIO: Montgomery, May (UMSP). 
PENNSYLVANIA: Allegheny, June (CMNH). WIS- 
CONSIN: Burnett, June (MS). 


Aethes angulatana (Robinson) 
(Fig. 2a—d) 


Conchylis angulatana Robinson (1869:286, pl. 8, fig. 
81) ( Lectotype ° designated by Klots [1942], type 
#7442, “Penn.,” ND, FW 5.0 mm long, gen. prep. 
MGP932 [Fig. 2d], in ANSP). Fernald (1882:25), 
Klots (1942:416), Handfield et al. (1997:44). 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Phalonia angulatana; Dyar et al. (1903:487), Forbes 
1923:508), McDunnough (1939:60), Brower 
1983:50). 

Aethes angulatana (not Robinson 1869); Razowski 
1986:394, 1995:139, 1997:123). 

Aethes angulatana; Grehan et al. (1995:25), Covell 
1999:62). 

Phtheochroa angulatana; Poole and Gentili (1996:876), 
Nielsen (1998:10). 

Male and female exterior. Head cream colored, 
often suffused with darker scales; labial palpus 3x 
longer than horizontal eye diameter. Notum and tegu- 
lum cream and mixed buff, brown, or black; hind leg 
cream, tibial spurs not contrasting. FW (Fig. 2a) 
4.8-7.6 mm long (n = 53). Ground color cream, often 
suffused with darker scales; markings a mixture of 
buff, brown, and black scales, fasciae outlined inter- 
mittently with darker scales; basal one-fourth of costa 
dark, costa and anal margin outlined with short striae; 
basal fascia extending to costa, protruding slightly into 
discal cell at midpoint; IM fascia extending into discal 
cell, often coalescing with discal spot, scattered scales 
angling from discal spot toward termen and tornus; 
anal crescent and OM fascia present; ST line extend- 
ing from tornus to apex; fringe of ground color, often 
bicolored, with darker basal band the color of FW 
markings. HW dark drab, fringe paler with darker 
basal band. 

Male genitalia (Fig. 2b, c) (n = 25). Valva broad, 
costa subsinuate, posterior margin broad, curved, a 
horseshoe-shaped row of dentiform spines on harpe; 
sacculus pointed. Median process of transtilla thumb- 
shaped, apex rounded, with a small terminal process. 
Aedeagus moderately thick, vesica bearing one long 
cornutus and a small irregularly shaped Thumncle of 
smaller cornuti. Anellus lobes absent. 

Female genitalia (Fig. 2d) (n = 23). Lamella post- 
vaginalis a small medial round process; bases of 
apophyses broad, posterior apophyses as long as ante- 
rior; antrum robust, long, funnel shaped, tapering an- 
teriorly; ductus bursae sclerotized, coalesced with cor- 
pus bursae, corpus bursae slanted to right when 
viewed as in Fig. 2d. Cervix bursae to the left and dor- 
sad of antrum when similarly viewed. 

Biology. Larval foodplants unknown. Adults col- 
lected mid-June to mid-September. Life cycle proba- 
bly univoltine. 

Diagnosis. A. angulatana is highly variable in size 
and markings, with some phenotypes resembling other 
species; it can be diagnosed best by genitalia. Its large 
lamella postvaginalis, and lack of a distal aedeagal 
sclerite differentiate it from A. promptana (Figs. 2d, 
1d, 2c, 1c); its narrower phallobase apex, and lack of 


VOLUME 56, NUMBER 4 


ventral cervix bursae differentiate it from from A. sex- 
dentata (Figs. 2d, 5d, 2c, 5c); its smaller lamella post- 
vaginalis, and absence of a blade-shaped aedeagal ca- 
rina differentiate it from A razowskii (Figs. 2d, 6d, 2c, 
6c); its slightly narrower IM fascia, and lack of a distal 
aedeagal sclerite also differentiate it from A razowskii 
(Figs. 2c, 7c, 2a,7a); and its broader harpe and less pro- 
nounced cervix bursae differentiate it from A. matheri 
(Figs. 2d, 8d, 2b, 8b). Melanics also occur. 

Discussion. This species was misidentified by Ra- 
zowski (1986, 1995, 1997) who based his identification 
on misidentified specimens. He did not report exam- 
ining the lectotype. Razowski’s A. angulatana is de- 
scribed later here as A. sexdentata, new species. 

Specimens examined (n = 53, including type). 
Counties and months of capture by State or Province: 
Canada. QUEBEC: Gatineau, July (UMSP). USA. 
ILLINOIS: Putnam, September (USNM). INDIANA: 
St. Joseph, July-September (UMSP). MAINE: 
Franklin, June; Washington, August (MAR). MICHI- 
GAN: Allegan, August; Cass, July, September; 
Cheboygan, July; Clinton, July, September; Kalama- 
zoo, September; Otsego, August; St. Clair, September; 
St. Joseph, September (BGS, GJB, JHW, MSUC, 
UMSP, WPW). MINNESOTA: Wilkin, July (UMSP). 
NEW JERSEY: Burlington, June, September, October 
(ANSP). PENNSYLVANIA: Allegheny, August, Sep- 
tember; Beaver, August, September (CMNH, 
USNM). VERMONT: Bennington, August; Chitten- 
den, August; Grand Isle, September (FMPS, MS). 
WISCONSIN: Burnett, August (MS). 


Aethes argentilimitana (Robinson) 


(Fig. 3a—d) 


Conchylis argentilimitana Robinson (1869:287, pl. 8, 
fig. 82) (Lectotype ¢ designated by Klots [1942], type 
#7441, “Penn.,” ND, gen. prep. MGP931 [Fig. 3c], 
FWs missing, in ANSP), Fernald (1882:25), Klots 
(1942:417), Handfield et al. (1997:44), Covell 
(1999:62). 

Conchylis labeculana Robinson (1869:287, pl. 8, fig. 
83) (Lectotype ¢ designated by Klots [1942], type 
#7443, “Penn.,” ND, gen. prep. MGP934 [Fig. 3b], 
FW 5.5 mm long [Fig. 3a], in ANSP), Fernald 
(1882:25), Klots (1942:417), new synonym. 

Phalonia argentilimitana; Dyar et al. (1903:487), Mc- 
Dunnough (1939:60), Brower (1983:50). 

Phalonia labeculana; Dyar et al. (1903:487), McDun- 
nough (1939:60). 

Aethes argentilimitana; Grehan et al. (1995:25), Ra- 
zowski (1997:123, figs. 51-53, 149). 

Phtheochroa argentlimitana; Poole and Gentili 
(1996:876). 


Phtheochroa labeculana; Poole and Gentili (1996:876). 
Aethes labeculana (not Robinson 1869); Razowski 

(1997:122). 

Male and Female exterior. Head, notum, and 
tegulum white; labial palpus 2x longer than horizontal 
eye diameter. Thoracic tuft inconspicuous, middle and 
hind legs predominantly buff, tibial spurs not contrast- 
ing. FW (Fig. 3a) 3.9-6.4 mm long (n = 81). Ground 
color white; markings buff to raw umber; buff and um- 
ber striae on costa and anal margin; basal fascia ex- 
tending to costa; often suffused basally with buff 
scales; IM fascia extending into discal cell, scattered 
scales reaching anal crescent; discal spot minute; OM 
fascia short, narrow, sometimes coalesced with ST line; 
ST line irregular, consisting of scattered scales reach- 
ing apex; fringe of ground color. HW drab; fringe pale, 
basal line present. 

Male genitalia (Fig. 3b, c) (n = 16). Costa of valva 
straight, posterior margin narrowly rounded, apex 
oblique, harpe spined; sacculus lobed. Aedeagus thick, 
vesica bearing one stout cornutus and a scobinate 
patch, lacking small cornuti. Anellus lobes absent. 

Female genitalia (Fig. 3d) (n = 11). Anterior and 
posterior apophyses subequal in length. Antrum fun- 
nel-shaped, apical half broad, shorter than in A. angu- 
latana, often twisted forward almost entire length of 
ductus bursae; ductus bursae scerlotized, coalesced 
with the elongate corpus bursae. Cervix bursae located 
at posterior end of corpus bursae, subequal in size to it. 

Diagnosis. A. argentilimitana is readily recognized 
by the golden buff FW markings. As noted by Razowski 
(1997), FW marginal striae are absent in some speci- 
mens, which makes their FW ground color solid white. 

Discussion. Although the type of A. argentilimi- 
tana lacks forewings, its genitalia are intact, and geni- 
talia of the type of A. labeculana do not differ from 
them. Razowski (1997) misidentifed A. labeculana, 
basing his identification on a misidentified specimen. 
He did not report examining the type. 

Biology. Larval foodplants unknown. May—October 
adult capture dates suggest the life cycle is multivol- 
tine. Sabourin has captured adults in dry, open areas of 
meadows and fields. 

Specimens examined (n = 81, including types of 
A. argentilimitana and A. labeculana). Counties and 
months of capture by State or Province: Canada. ON- 
TARIO: Muskoka, July (ANSP). USA. ILLINOIS: 
Putnam, May, June; Union, July (INHS). INDIANA: 
Elkhart, May; LaGrange, August; Perry, June; St. 
Joseph, August (JV, UMSP). KENTUCKY: Barren, 
September; Meade, May; Rockcastle, May (ANSP). 
MAINE: Kennebec, July (MAR). MASSACHU- 
SETTS: Dukes, July, August (ANSP). MICHIGAN: 


bo 
NS) 
pS 


Allegan, May; Barry, August; Cass, June; Clinton, 
June; Washtenaw, July (JHW, UMSP, WPW). MIN- 
NESOTA: Anoka, May; Beltrami, June, July; Clearwa- 
ter, June-August; Marshall, August (UMSP). MISSIS- 
SIPPI: Lee, May; Oktibbeha, April, May; Tishomingo, 
June, July (BMMS, UMSP). NEW JERSEY: Burling- 
ton, June-August (ANSP). OHIO: Adams, July; Cler- 
mont, May; Hamilton, May-July, September, October 
(ANSP). PENNSYLVANIA: Allegheny, June, August; 
Bucks, July (AMNH, CMNH). VERMONT: Chitten- 
den, May, July; Grand Isle, August (MS, FMPS). WIS- 
CONSIN: Burnett, May, June, August (UMSP). 


Aethes interruptofasciata (Robinson), 
revised status 
(Fig. 4a—d) 


Conchylis interruptofasciata Robinson (1869:287, pl. 
8, fig. 85) (Lectotype d designated here, type #7444, 
“Penn.,” ND, gen. prep. MGP933, FW 5.0 mm 
long, in ANSP), Fernald (1882:25). 

Phalonia interruptofasciata; Dyar et al. (1903:487) 
Forbes (1923:506). 

Phalonia aureana Busck (1907:25) (Holotype °: Penn- 
sylvania, Oak Station, F. Marloff, gen. prep. USNM 
#24375, in USNM), Forbes (1923:507), McDun- 
nough (1939:60), Procter (1946:308), Covell (1999: 
62), Brown and Lewis (2000:1021). 

Phalonia sublepidana Kearfott (1907:82) (Lectotype ¢ 
designated by Klots [1942], N. J., Caldwell [Essex 
Co.], W. D. Kearfott, 8 July 1900, gen. prep. 
MGP810, in AMNH), Forbes (1923:507), Mc- 
Dunnough (1939:60), Procter (1946:308), Brower 
(1983:50), new synonym. 

Phalonia interruptofasciana; McDunnough (1939:60), 
Brower (1983:50). Misspelling. 

Phtheochroa aureana; Poole and Gentili (1996:876). 

Phtheochroa interruptofasciana; Poole and Gentili 
(1996:876). Misspelling. 

Phtheochroa sublepidana; Poole and Gentili (1996:876). 

Aethes labeculana (not Robinson 1869); Razowski 
(1997:122, figs. 49, 50, 147, 148). 

Male and female exterior. Head, tegulum, and 
thoracic tuft mixed cream and buff, remainder of no- 
tum predominantly fuscous; labial palpus 2.5x longer 
than horizontal eye diameter. Fore- and midlegs fus- 
cous anteriorly, buff posteriorly, mixed fuscous and 
buff at articulations, hindleg predominantly buff, some 
fuscous scales posteriorly. FW (Fig. 4a) 4.7-7.0 mm (n 
= 32). Ground color cream, suffused with cinnamon 
and fuscous scales; markings varying from cinnamon to 
burnt umber; basal fascia cinnamon; fuscous striae on 
costa and anal margin; IM fascia cinnamon, short, ex- 
tending barely into discal cell; anal crescent narrow, 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


umber, darker than IM fascia, or absent; fuscous scales 
along veins from discal cell to termen; terminal half of 
FW suffused with fuscous scales in sublepidana phe- 
notype; discal spot not prominent, some cinnamon 
scaling between IM and OM fasciae; OM fascia ex- 
tending to tornus, often paler below costa; ST line ab- 
sent; fringe cream; apex obtuse. HW dark drab; fringe 
a mixture of drab and paler scales. 

Male genitalia (Fig. 4b, c) (n = 10). Costa of valva 
subsinuate, posterior margin narrowly rounded, 
oblique, harpe unspined; sacculus lobed. Sicklelike 
apical structures of socii long. Aedeagus long, thin, 
curved, vesica with one long cornutus. Anellus lobes 
absent. Juxta small, rotund, a V-shaped incision at its 
junction with aedeagus. 

Female genitalia (Fig. 4d) (n = 5). Anterior and 
posterior apophyses subequal in length to papillae 
anales. Sterigma small, crescent shaped; lamella post- 
vaginalis a medial spot; antrum short, cylindrical, 1/6 
length of ductus bursae; ductus bursae long, posterior 
1/2 sclerotized. Cervix bursae a small fingerlike 
process on basal left margin of ductus bursae when 
viewed as in Fig. 4d. 

Diagnosis. Pale specimens of A. interruptofasciata 
can be differentiated from A. argentilimitana by their 
broader basal fascia, and the OM fascia extending to 
the tornus (Figs. 4a, 3a). The two taxa also differ 
markedly in genitalia; males of A. argentilimitana pos- 
sess a spined harpe and those of A. interrwptofasciata 
lack it (Figs. 4b, 3b). Material labeled as A. sublepi- 
dana suggests an undersized A. interruptofasciata; no 
consistent genitalic differences were found between 
them. 

Discussion. A. interruptofasciata is resurrected 
from the synonymy of A. labeculana. Razowski (1997) 
synonymized A. interruptofasciata under A. labecu- 
lana based on a misidentified specimen of A. labecu- 
lana which actually is A. argentilimitana. Variability in 
size and maculation of A. interruptofasciata has led to 
descriptions of some of its phenotypes as separate 
species. We designate a lectotype to prevent future 
confusion about the identity of this taxon. 

Biology. Larval foodplants unknown. Adults cap- 
tured May 20-August 4. Life cycle probably multivol- 
tine. Sabourin has captured adults in deciduous forest 
openings and blueberry thickets. 

Specimens examined (n = 32, including types of 
A. interruptofasciata, sublepidana, and aureana). 
Counties and months of capture by State or Province: 
Canada. MANITOBA (no county system): Aweme, 
June (CNC). NOVA SCOTIA: Queens, July (NSPM). 
USA. ILLINOIS: Putnam, July (USNM). MAINE: 
Washington, July (MAR), MICHIGAN: Cheboygan, 


VOLUME 56, NUMBER 4 


July (BGS). MISSOURI: Wayne, June (JRH). NEW 
JERSEY: Essex, July (AMNH). PENNSLYVANIA: Al- 
legheny, June-August (AMNH, CMNH). WEST VIR- 
GINIA: Kanawha, May (UMSP). WISCONSIN: Bur- 
nett, June, July; Douglas, July; Oneida, July; Vilas, 
June (EME, GJB, MS, UMSP). 


A. sexdentata Sabourin and Miller, new species 


(Fig. 5a—d) 


Aethes sp. nr. angulatana; Grehan et al. (1995:25). 
Aethes angulatana (not Robinson, 1869); Razowski 

(1997:123, figs. 54, 55, in part). 

Male and female exterior. Head and notum 
white, dusted with buff; labial palpus 3x longer than 
horizontal eye diameter. Tegulum mixed raw umber, 
clay, and buff; midleg fuscous anteriorly, with buff 
scales at junctions of tarsomeres, buff posteriorly; 
hindleg predominately buff, some fuscous scales on fe- 
mur, tibial spurs fuscous anteriorly, the shorter of the 
pairs usually darker. FW (Fig. 5a) 5.7-8.2 mm long (n 
= 83). Ground color white, dusted with buff scales ex- 
cept along costa; costa and anal margin outlined with 
short striae; markings raw umber, clay, and buff; fas- 
ciae black bordered; basal fascia broad, extending to 
costa; IM fascia extending into discal cell, coalesceing 
with discal spot; scattered scales from discal spot an- 
gled toward tornus and beyond apex of anal crescent; 
OM fascia a broad spot; ST line extending from tornus 
to apex; fringe buff with less mixture of variously col- 
ored scales than in A. angulatana; terminal line darker 
than fringe. HW dark drab; fringe cream with darker 
basal band. 

Male genitalia (Fig. 5b, c) (n = 37). Costa of valva 
slightly curved, posterior margin broad, rounded, 4-12 
spines on harpe, but usually 6; sacculus rounded. Me- 
dian process of transtilla subtriangular. Aedeagus 
thick, vesica bearing one large cornutus and a bundle 
of smaller cormuti. Phallobase broadening at apex. 
Anellus lobes absent. 

Female genitalia (Fig. 5d) (n = 19). Anterior and 
posterior apophyses subequal in length, longer than 
papillae anales. Lamella postvaginalis a small medial 
plate; terminal half of antrum broadening sinistrad 
when viewed as in Fig. 5d; cervix bursae circular, pro- 
duced ventrad to base of antrum; tergum 7 lightly 
sclerotized. 

Types. Holotype: VERMONT, Grand Isle Co., 
Grand Isle, Lovers Lane, 4, 07/27/1995 (M. S. Griggs), 
FW 7.0 mm long, gen. prep. MS01046 (Fig. 5b, c) 
(UMSP). Paratypes (n = 78): Canada. NOVA SCO- 
TIA: Colchester Co., Debert, 3, 07/20/1961 (D. C. Fer- 
guson), gen. prep. MS00308 (NSPM). ONTARIO: 
Grand Bend [Huron Co.], 2°, 07/06/1939 (T. N. 


Freeman), gen. prep. MS97249. Pt. Colborne [Welland 
Co.], d, 07/13/1932, gen. prep. Ph23; 4, 05/07/1933 (J. J. 
deGryse), gen. prep. MS97308 (CNC). Trenton 
[Northumberland Co.], 2, 07/12/1912 (Evans) (CNC). 
QUEBEC: Montreal [Laval Co.], 3, 07/01/1985 (B. 
Landry) (MS). USA. CONNECTICUT: Washington 
[Litchfield Co.], AL, J, 06/21/1960, gen. prep. MS00407, 
coll. #63404; 3, 06/22/1960, coll. #63404; 3, 06/20/1961, 
gen. prep. MS97292, coll. #63406; J, 07/17/1960, gen. 
prep. MS01063, coll. #63399: d, 07/21/1960, gen. prep. 
MS00407, coll. # 63400; 3, 06/25/1962 (S. A. Hessel), 
gen. prep. MS97260, coll. # 63421(YPM). Windham 
Co., Putnam, 3, 07/14/1959 (Klots), gen. prep. 
MS01028 (AMNH). ILLINOIS: Algonquin [McHenry 
Co.], 3, 07/07/1909, gen. prep. 00738IG. Clark Co., 
Rocky Branch Preserve, UVL, 2, 07/03/1995 (T. Harri- 
son), gen. prep. MS98427 (INHS). INDIANA: Elk{hart] 
Co., LT, d, 06/21/1997, gen. prep. MS98413. St Joseph 
Co., LT, 4, 07/31/1996 (J. Vargo), gen. prep. MS97290 
(UMSP). MAINE: Steuben [Washington Co.], 4, 
(6/11/1999, gen. prep. 01072IG; °, 07/17/1999, gen. 
prep. MS01073; d, 06/26/2000 (M. A. Roberts), gen. 
prep. MSO1066 (MAR). MASSACHUSETTS: Lan- 
caster [Worcester Co.], 3, 07/15/1993 (E. Peters) (MS). 
MICHIGAN: Cass Co., Westrate farm, LIT, 4, 
07/16/1999 (J. Vargo), gen. prep. MS99438 (UMSP); 
T5S R14W Sec. 31, 4, 06/11/1987 (W. P. Westrate), gen. 
prep. MS01051 (WPW). Ingham Co., T4N R2W Sec. 
35, 3 6, 07/20/1968 (J. P. Donahue), gen. prep. 01105IG. 
Lenawee Co., T8S R2E Sec. 31, 3 4, 07/08/1969. Liv- 
ingston Co., George Reserve, °, 07/19/1934 (W. C. Stin- 
son), gen. prep. MS00502. Midland Co., 2, 08/09/1958 
gen. prep. MSO01093; d, 08/11/1958 (R. R. Dreisbach), 
gen. prep. MS00727 (MSUC). Muskegon Co., 
Muskegon S|tate] Park] dunes, 2, 08/11/1989 (G. 
Balogh), gen. prep. 97293 (GJB). Oakland Co., 2, 
06/26/1933 (W. C. Stinson), gen. prep. 010941G 
(MSUC). MINNESOTA: Itasca Pk., LaSalle Valley 
[Clearwater Co.], AL, °, 07/09/1940 (C. E. Mickel), gen. 
prep. MSO01053. Ramsey Co., North Oaks, 3, 06/28/1965 
gen. prep. JAB122; d, 08/02/1965 (W. E. Miller), gen. 
prep. JAB121. Wilkin Co., ¢, 07/11/1937 (D. G. Den- 
ning), gen. prep. 011031G. MISSISSIPPI: Claiborne 
Co., Rocky Springs, 3, 05/10/1970, gen. prep. MS00058, 
coll. #35670. Hinds Co., Clinton, ¢, 11/13/1958, gen. 
prep. MS99495, coll. #287; 3, 05/07/1971 (B. Mather), 
gen. prep. MS99172, coll. #35561 (UMSP). NE- 
BRASKA: Cherry Co., Valentine N[atl.]| Wildlife] 
Rlefuge], Hackberry Lk, LT, 6, 06/21/1983; 4, 
06/28/1983: 3, 06/29/1983, gen, prep. MS01223; 9, 
06/30/1983 (R. W. Hodges) (USNM). NEW JERSEY: 
New Lisbon [Burlington Co.], 4, 06/17/1933; 4, 
06/18/1933 (E. P. Darlington), gen. prep. MS00243. 


PENNSYLVANIA: Philadelphia [Philadelphia Co.], ¢, 
07/05/1914 (ANSP). VERMONT: Chittenden Co., 
Colchester, 07/04/1993, 2, 3d, gen. prep. MS01037; °, 2 
3, 07/11/1993, gen. preps. MS01150, MS01148; Col- 
chester, railway by bog, 2 6, 08/05/1992, gen. prep. 
MS95013: Shelburne, 4, 07/04/1993: S. Burlington, Q, 
07/29/1998, gen. prep. MS01146 (MS); 2, 07/12/1992, 
gen. prep. MS97250; 3, 07/28/1992; 3 (Fig. 5a), 
07/07/1993 (M. Sabourin), gen. prep. MS97207. Grand 
Isle Co., Grand Isle, Lovers Lane, 2, 07/27/1995 (M. S. 
Griggs), gen. prep. MS00056 (UMSP). WASHING- 
TON: Pullman [Whitman Co.], 3, 05/1935, reared ex 
Solidago stalks, gen. prep. MS00060 (EME); 3 d, 2 ° 
05/1935 (J. F. G. Clarke), reared ex Solidago stalks, gen. 
preps. USNM23877, 23878, 23880, 23980, wing slides 
MS23877, USNM23878. Walla Walla [Walla Walla Co.], 
3, 9, 06/06/1931(D. R. Brannon), gen. preps. 
USNM23879, 23981(USNM). WEST VIRGINIA: 2, no 
data, gen. prep. MS00013, pseudotype argentilimitana 
(Klots 1942), Grote and Robinson type #23031(AMNH). 
WISCONSIN: Burnett < Grantsburg, AL, 4, 
06/07/2000; 3, 06/26/2000; 2°, 06/23/2001, gen prep. 
MS01482; 2, 07/01/2000, een. prep. MS00602; 2°, 
07/09/2000, gen. prep. MS00621, wing _ slide 
MS00621W; 6, 08/09/2000; T40N R1SW Sec. 23, LT, 6, 
06/30/2001, gen. prep. MS01494; 3, 07/14/2001 (M. 
Sabourin), gen. prep. MSO0601 (MS, ZMUH). Dane 
Co., Nevin Marsh, ¢, 06/26/1974 (D. T. Bach) (UWEM). 

Diagnosis. A. sexdentata most resembles A. fernal- 
dana (Walsingham 1879), the wings of which were il- 
lustrated by Walsingham (1879), and the male geni- 

talia by Razowski (1964). We examined the A. 
fernaldana syntypes enumerated below. 

A. sexdentata differs in its white FW ground color, 
and markings predominantly of raw umber and buff 
with black borders, versus the yellow ground with 
ochreous markings in A. fernaldana. In addition, the 
OM fascia and ST line in A. sexdentata are conspicu- 
ous and concolorous with the IM fascia, whereas in A. 
fernaldana the OM fascia and ST line are paler than 
the IM fascia and hardly differentiated from the 
ground color. 

In male genitalia, A. sexdentata differs in its more 
sinuate ventral margin of the valva, and more rounded 
ventral margin of the sacculus. Further, although FW 
length of A. fernaldana ranges within that given above 
for A. sexdentata, the single large cornutus of A. sex- 
dentata is twice as thick and nearly twice as long as 
that of A. fernaldana, and the small cornuti of A. sex- 
dantata are larger and twice as numerous as those of 
A. fernaldana. The genitalia preparation of the one fe- 
male syntype of A. fernaldana was in too poor condi- 
tion for comparison. The cervix bursae of A. sexdentata 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


is ventrad to the antrum, which otherwise separates fe- 
males of the new species from all known Aethes. 

A. fernaldana specimens examined (n = 3). FW 
length 7.0-8.5 mm. Type H. T., Hatchet Creek, Shasta 
Co., California, 07/14-17/1871, Cochylis fernaldana 
Wlsm., type ¢ figd. & descr... ., B. M. ¢ genitalia slide 
No. 4777. Same data except Cochylis fernaldana 
paratype, B. M. d genitalia slide No. 12950. Same data 
except B. M. 2 genitalia slide No. 18564 (BM). So far 
as we know, no types have been formally designated, 
so these specimens are considered syntypes here. 

Discussion. This is Razowskis A. angulatana, the 
identity of which was based on a misidentified specimen. 
Females were associated by FW color pattern and si- 
multaneous capture. One melanic specimen was found. 
One of the most widely distributed Nearctic Aethes, A. 
sexdentata ranges from Nova Scotia west to Washing- 
ton State, and south to Mississippi and Nebraska. 

Etymology. The name sexdentata refers to the 
spines on the harpe. 

Biology. Solidago sp. (Asteraceae) is a larval food- 
plant (Razowski 1997, and data on pinned specimens). 
Adults were captured May 7—August 11, suggesting a 
univoltine life cycle. 


Aethes razowskii Sabourin and Miller, 
new species 


(Fig. 6a—d) 


Phalonia promptana (not Robinson, 1869); Forbes 
(1923:508, in part), Brower (1983:50, in part). 
Aethes promptana (not Robinson, 1869); Grehan et al. 
(1995:25), Razowski (1997:124, figs. 56-60, 150-152). 
Male and female exterior. Head, notum, and 
tegulum cream colored; labial palpus 3x longer than 
horizontal eye diameter. Fore- and midlegs darker an- 
teriorly than posteriorly, hindleg with some brownish 
scales on femur, the smaller tibial spurs darker than 
the larger ones. FW (Fig. 6a) 4.7-8.6 mm long (n = 
34). Ground cream colored; markings buff, orange yel- 
low, dark brown, and fuscous; all fasciae bordered by 
darker scales; basal fascia narrow, extending to costa; 
striae on costa and anal margin not conspicuous; IM 
fascia narrow, extending into cell, coalescing with dis- 
cal spot; discal spot paler than IM fascia, often barely 
contrasting with ground color; anal crescent present; 
OM fascia short; ST line extending to vein R., hardly 
differentiated from ground, orange yellow with a few 
umber specks to more contrasting and brown in some 
females; fringe of ground color. HW dark drab; fringe 
paler with narrow basal band. 
Male genitalia (Fig. 6b, c) (n = 23). Costa of valva 
subsinuate, posterior margin broad, dorsally oblique, 
U-shaped row of spines on harpe, spines on exterior 


VOLUME 56, NUMBER 4 


margin longer than those on interior; sacculus broadly 
rounded. Aedeagus moderately thick, vesica bearing a 
blade-shaped beak or carina, an elongate scobinate 
patch, and one large, thick cornutus. Anellus lobes ab- 
sent. 

Female genitalia (Fig. 6d) (n = 4). Anterior and 
posterior apophyses subequal in length to papillae 
anales. Lamella postvaginalis a subquadrate medial 
plate; antrum cylindrical; ductus bursae sclerotized en- 
tire length. Cervix bursae subquadrate, not notably 
separated from corpus bursae. 

Types. Holotype: MICHIGAN, Schoolcraft Co., 
T42N RI6W Secs. 11, 13, 5, 07/04/1987 (G. Balogh), 
FW 6.5 mm long, gen. prep. MS01050 (Fig. 6b) 
(AMNH). Paratypes (n = 19): Canada. NOVA SCO- 
TIA: Round Hill [Annapolis Co.], 6, 06/24/1961 (H. 
Stultz) (NSPM). QUEBEC: Knowlton [Brome Co.], 2 
3, 06/09/1936 (G. S. Walley), gen. prep. Ph13 (CNC). 
USA. ALABAMA: Winston Co., d, 04/11/1999, gen. 
prep. MS99196; °, gen. prep. MS99197 (MS). CON- 
NECTICUT: Windham Co., Putnam, °, 08/13—16/1961 
(A. B. Klots), gen. prep. MS00202 (AMNH). IDAHO: 
Wallace [Shoshone Co.], 3, 3000 ft., Sweadner collec- 
tion, 06/22-30, gen. prep. MS00245 (CMNH). INDI- 
ANA: St. Joseph Co., LT, d, 06/20/1999 (J. Vargo), gen. 
prep. IG (JV). MAINE: Rangeley [Franklin Co.], }, 
07/10/1938; 2, 06/23/1938 (V. H. dos Passos), gen. preps. 
00062IG, MS00016 (AMNH). MARYLAND: Balti- 
more Co., Prettyboy Reservoir, 3, 08/24/2001, coll. #4, 
gen. prep. MS02029; Soldiers Delight, 6, 04/25/1998; 3, 
05/25/1997, gen. prep. MS02205; 3, 05/15/2000, gen. 
prep. MS0220; 3, 06/14/1997: 2 3, 07/05/1997; 4, 
08/07/1997, 3, 09/06/1998; 2, 09/30/1998, gen. prep. 
MS02206; ¢, 09/19/1998, gen. prep. MS02182; 3, 
05/15/1998, gen. prep. IG02184 (J. Glaser) (JDG). 
MICHIGAN: Allegan Co., T2N R14W Sec. 26, (Fig. 
6a), 5/30/1986 (G. Balogh), gen. prep. MS97310 (GJB). 
Cass Co., T5S R14W Sec. 31, 4, 07/25/1996 (W. P. Wes- 
trate), gen. prep. IG (WPW); LT, 2 4, 09/03/1999, gen. 
prep. IG; 4, 09/25/1999 (J. Vargo), gen. prep. MS99529 
(UMSP). Gladwin Co., 3, 06/25/1959 (R. R. Dreisbach), 
gen. prep. 01108IG (MSUC). Livingston Co., George 
Reserve, AL, 4, 07/22/1938 (S. Moore), gen. prep. 
MS01101 (UMMZ). Schoolcraft Co., T42N RI6W 
Secs. 11, 13, 4, 07/04/1987 (G. Balogh), gen. prep. 
MS01050 (GJB). St. Joseph Co., Three Rivers, LT, 3, 
05/29/1999 (J. Vargo), gen. prep. MS99439, wing slide 
MS99439W (UMSP). Washtenaw Co., T3S R3E Sec. 
29, 3, 05/30/1992 (B. G. Scholtens), gen. prep. 000271G 
(BGS); T2S R3E Sec. 6, 2, 07/19/1993 (J. H. Wilter- 
ding), gen. prep. MS00499 (JHW). NEW HAMP- 
SHIRE: Coos Co., Whitefield, BLT, 2, 07/11/1992 (W. 
Kiel), gen. prep. MS01496 (MS). VERMONT: Ferdi- 


22. 


nand [Essex Co.], 3, 06/23/1993 (M. Sabourin), gen. 
prep. 002471G (UMSP). 

Diagnosis. A. razowskii most resembles A. promp- 
tana. FW markings in A. razowskii are usually more 
contrasting with ground color and outlined by darker 
scales (Figs. 6a, 1a). In male genitalia, A. razowskii dif- 
fers from A. promptana in its oblique posterior margin 
of the valva and well rounded ventral margin of the 
sacculus, compared with the curved posterior margin 
of the valva and straight margin of the sacculus in A. 
promptana (Figs. 6b, 1b). Also, the aedeagus of A. ra- 
zowskii has a blade-shaped carina, which is absent in 
A. promptana (Figs. 6c, 1c). In female genitalia, the 
antrum of A. razowskii is more cylindrical and the 
lamella postvaginalis smaller than in A. promptana 
(Figs. 6d, 1d). A. razowskii is superficially similar to 
some individuals of A. angulatana, but in A. razowskii 
the IM fascia is narrower, the ST line paler, and the 
genitalia are different (Figs. 6a—d, 2a—d). 

Discussion. Razowski (1997) and earlier authors 
misidentified this species as A. promptana. Females 
were associated by FW color pattern and simultaneous 
capture. 

Etymology. This species is named in honor of Dr. 
Josef Razowski in recognition of his continuing efforts 
to illuminate the world tortricid fauna. 

Biology. Larval foodplants are unknown. May—Sep- 
tember adult capture dates suggest a multivoltine life 
cycle. 


Aethes westratei Sabourin and Miller, 
new species 


(Fig. 7a—d) 


Male and female exterior. Head and notum usu- 
ally unicolorous cream to tawny, antennal scape darker 
than head; labial palpus white apically, 3x longer than 
horizontal eye diameter. Thoracic tuft conspicuous, 
tegulum brown, darker than notum; fore- and midlegs 
fuscous anteriorly with buff scales at articulations, 
hindleg predominately buff except for fuscous scales 
anteriorly on shorter tibial spurs. FW (Fig. 7a) 6.9 mm 
long (n = 9). Ground color cream, suffused with 
brownish scales, basal area suffused with tawny scales, 
posterior 3/4 often suffused with grayish scales; mark- 
ings brown; fasciae with darker brown or black bor- 
ders; basal fascia extending slightly into discal cell at 
midpoint, also to costa; costa and anal margins with 
short striae; IM fascia broad, coalescing with discal 
spot; anal crescent and OM fascia present; ST line ir- 
regular, but extending to apex; fringe cream with some 
brown scales; terminal line darker than fringe. HW 
olive-brown; fringe paler, with basal band. 

Male genitalia (Fig. 7b, c) (n = 4). Costa of valva 


bo 
i) 
(Oa) 


straight, posterior margin rounded, sinuate, harpe 
spined; sacculus subtriangular, ventrally rounded. Me- 
dian process of transtilla subtriangular. Aedeagus 
thick, vesica bearing one large cornutus, a bundle of 
small cornuti, and a distal crescent-shaped sclerite. 
Anellus lobes absent. 

Female genitalia (Fig. 7d) (n = 1). Anterior and 
posterior apophyses equal in length; antrum funnel 
shaped, almost as long as ductus bursae, right margin 
straight, left margin bowed at 1/3 its length when 
viewed as in Fig. 7d. Cervix bursae sinistrocaudad of 
corpus bursae when similar ly viewed, and small. Cor- 
pus bursae elongate, slightly slanted. 

Types. Holotype: MICHIGAN, Cass Co., T5S 
RI4W Sec. 31, UV, d, 09/02/1995 (M. C. Nielsen), FW 
6.5 mm long, gen. prep. MS97289 (AMNH). 
Paratypes (n = 8): MICHIGAN: Barry Co., Shaw Lk., 
T3N RLOW Sec 3, 2 d, 09/09/1986 (G. Balogh) (GJB). 
Cass Co., LT, 5, 09/06/1997, gen. prep. MS97325; ° 
(Fig. 7a), same data (J. Vargo), gen. prep. MS98146 
(UMSP): T5S R14W Sec. 31, UV, d, 09/02/1995 (M. C. 
ee gen. are. MS0i039 (MSUC); same locality, 

, 09/02/1988: 3, 09/09/1988 (W. P. Westrate), gen. 
prep. M01048, wing slide MSO1048W (WPW); Dr. 
Lawless Clounty] P[ark], T6S R13W Sec. 32, UV, d, 
09/01/1995 (M. C. Nielsen) (MSUC). 

Diagnosis. A. westratei most resembles A. angula- 
tana superficially and genitalically. The broad IM fas- 
cia is a distinguishing feature, being broader in A. wes- 
tratei than in A. angulatana (Figs. 7a, 2a). In male 
genitalia, the harpe of A. westratei is more heavily 
spined than in A. angulatana, the posterior margin of 
the valva less sinuous, and the sacculus less acute 
(Figs. 7b, 2b). In female genitalia, the anterior and 
posterior apophyses are shorter and the corpus bursae 
less slanted than in A. angulatana (Figs. 7d, 2d). 

Discussion. Females were associated by FW color 
pattern and simultaneous capture. 

Etymology. This species is named in honor of 
William P. Westrate in recognition of his activities to il- 
luminate southern Michigan’s flora and fauna. 

Biology. Larval foodplants unknown. September 
adult capture dates suggest a univoltine life cycle. 


Aethes matheri Sabourin & Miller, 
new species 
(Fig. 8a—d) 


Phalonia angulatana (not Robinson, 1869): Kimball 

(1965:270). 

Male and female exterior. Head, notum, and 
tegulum usually cream or buff; labial palpus 2.5x 
longer than horizontal eye diameter. Hindleg without 
markings. FW (Fig. 8a) 4.4-6.6 mm long (n = 57). 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Ground color variable, usually cream or buff; markings 
buff and brown or cinnamon, fasciae outlined with fus- 
cous scales; basal 1/4 of costal margin fuscous; short 
striae on costal and anal margins; basal fascia variably 
projecting slightly into discal cell at midpoint and ex- 
tending to costa or costal half, diffuse, or absent; IM 
fascia extending into cell, bordered by or joining discal 
spot; anal crescent and OM fascia present; ST line 
present, not reaching apex, variable in color, often not 
differentiated; terminal line darker than fringe; fringe 
a mix of cream, cinnamon, buff, and brown scales, 
paler at tornus. HW pale drab; fringe paler with thin 
basal band. 

Male genitalia (Fig. 8b, c) (n = 34). Costa of valva 
straight, posterior margin narrowly rounded, harpe 
spined; sacculus pointed. Median process of transtilla 
triangular, rounded apically. Aedeagus laterally trian- 
gular at apex, vesica bearing one stout cornutus and a 
eral of smaller cornuti. Anellus lobes absent. 

Female genitalia (Fig. 8d) (n = 17). Posterior 
apophyses longer than anterior; left margin of antrum 
irregular when veiwed as in Fig. 8d; ductus bursae co- 
alescing with corpus bursae; cervix bursae sinistrocau- 
dad of corpus bursae when viewed as in Fig. 8d, sub- 
quadrate; corpus bursae elongate; tergum 7 lightly 
sclerotized. 

Types. Holotype (Fig. 8a): MISSISSIPPI, Long 
Beach [Harrison Co.], 3, 03/30/1997 (R. Kergosien), 
gen. prep. MS99483, FW 5.0 mm long, coll. #180525 
(AMNH). ). Paratypes ( n = 56): FLORIDA: Dade Co., 
Florida City, °, 04/02/1947; d, gen. prep. MS01137; 3 
05/02/1947, gen. prep. MSO1134; ¢, 05/04/1047, gen 
prep. MSO1L109: , 05/06/1947 (O. Buchholz), gen. 
prep. MSO1101; R. Palm Pk., AL, 3 2, 03/14/1938 (E. P. 
Darlington), gen. preps. 01136, MS01089 (ANSP). 
ILLINOIS: Coles Co., T12N R6E Sec. 11, UVL, 2, 
06/05/1997 (T. Harrison), gen. prep. MS98437. Putnam 
Co., 3, 07/06/1967 (M. O. Glenn), gen. prep. MS97297 
CONPEIS) 5 IUNIDIUNINUNS lalikirera]| (Co Ibi 3 6. 
05/29/1999, gen. prep. MSO1080. a . Joseph Co, ILI, 6. 
(6/20/1999, gen. prep. MSO06L1: 4 5, 06/23/1999, gen. 
prep. MS99508: « 3, 08/07/1999 (J. Vargo), gen. prep. 
002541G (UMSP). MAINE: Bangor iPenckeeot (Co. |, 
overgrown field across from Roadway Inn, Cornus, 
Alder, Spiraea, common, 08:50-09:50 h, 2 4, 
06/20/1997 (M. Sabourin), gen. prep. OO707IG (MS); d, 
same data, gen. prep. MS97350 (UMSP). Steuben 
[Washington Co.], 3, 06/10/1991, gen. prep. G1797; 4, 
07/07/1993 (M. A. Roberts), gen. prep. G1791 (MAR). 
MARYLAND: Baltimore Co.,Towson, ¢, 06/05/1999 (J. 
Glaser), gen, prep. 02204IG (JDG). MICHIGAN: 
Barry Co., Shaw Lk., T3N R1OW Sec. 3, d, 06/30/1989 
(G. Balogh), gen. prep. MS99189 (GJB). Shiawassee 


VOLUME 56, NUMBER 4 


Co., Moon Lk., T5N RIE Sec. 21, d, 06/25/1969 (J. P. 
Donahue), gen. prep. MS01100 (MSUC). St. Joseph 
Co., Three Rivers, LT, 2 4, 05/9/1999, gen. preps. 
MS99226, 99447; °, same date (J. Vargo), gen. prep. 
MS99227 (UMSP). MISSISSIPPI: Hancock Co., Bay 
St. Louis, °, 03/24/1972 (R. Kergosien), gen. prep. 
MS99170, coll. #106143. Harrison Co., Handsboro, 2, 
04/14/1966 (R. T. Taylor), gen. prep. IG, coll. #21264; 
Long Beach, °, 04/14/1978, gen. prep. MS01132, coll. 
#126347; J, 03/31/1997, gen. prep. IG, coll. #180569; 
d, 04/09/1997, gen. prep. IG, coll. #180549; d, 9, 
06/29/1997 (R. Kergosien), coll. #180550, #180570, 
gen. prep. IG (UMSP). Hinds Co., Clinton, ¢, 
09/02/1962 (B. Mather), coll. #11986 (BMMS): ¢, 
03/06/1995 (M. & E. Roshore), coll. #180543; Jack- 
son, 3, 04/07/1963, gen. prep. MS01082, coll. #13179; 
lOO MIoIs cents prep» IG, Weollk 438303:5 o 
04/30/1966, gen. prep. MSO01131, coll. #21278; 2, 
10/08/1966 (B. Mather), gen. prep. MS99436, coll. 
#22976. Jackson Co., Shepard S[tate] Plark], 4, 
08/28/1965, gen. prep. MS99040, coll. #20421; d, 
08/15-22/1995, gen. prep. 99524, coll. #175647; 2, 
09/6-11/1995, gen. prep. MS99275, wing slide 
MS99275W, coll. #164312. Rankin Co., Pearl, 9°, 
03/31/1963 (B. Mather), gen. prep. IG, coll. #13177 
(UMSP). Tishomingo Co., J. P. Coleman S[tate] P[ark], 
3, 06/10-24/1995, gen. prep. MS99282, coll. #164313 
(BMMS): d, 07/10-21/1995 (R. Kergosien), gen. prep. 
99485, coll. #180567 (UMSP). MISSOURI: Columbia 
[Boone Co.], 2, 07/22/1985 (W. S. Craig), gen. prep. 
MS99253 (UMRM). NORTH CAROLINA: Raleigh 
[Wake Co.], 3, 08/10/1938 (M. W. Wing), gen. prep. 
MS01135 (ANSP). PENNSYLVANIA: Allegheny Co., 
4 km N of Tarentum, 4, 06/22/1989 (W. Zanol), gen. 
prep. MS01107; Pittsburgh, 2, 06/21/1905 (H. Engel), 
gen. prep. 01095; 2, 06/27/1907 (H. Kahl), gen. prep. 
MS00501 (CMNH). TENNESSEE: Oak Ridge 
[Roane Co.], 3, 04/27/1966 (B. Mather), gen. prep. 
MS99441, coll. #21217 (UMSP). TEXAS: Beaumont 
[Jefferson Co.], 2, 08/02/1991 (C. Bordelon), gen. prep. 
MS01149 (MS). 

Diagnosis. A. matheri most resembles A. angula- 
tana. Superficially, A. matheri differs in its smaller size, 
more obscure basal fascia, narrower IM fascia, and ST 
line not reaching apex (Figs. 8a, 2a). In male genitalia 
of A. matheri, the valva is narrower distally, the base of 
the transtilla median process broader, and the apex of 
the aedeagus more acute (Figs. 8b, c, 2b, c). In A. 
matheri female genitalia, the left side of the antrum is 
more irregular, cervix bursae more conspicuous, and 
corpus bursae straighter than in A. angulatana when 
viewed as in Figs. 8d and 2d. A. matheri differs from 
A. sexdentata in male genitalia, the former, for ex- 


229 


ample, having an acute sacculus, and the latter an ob- 
tuse one (Figs. 8b, 5b). 

Discussion. The triangular process of the aedeagus 
can be seen by brushing away scales at the tip of the 
abdomen. Northern specimens of A. matheri are 
larger and have a less acute termination of the aedea- 
gus. Females were associated by FW color pattern and 
simultaneous capture. 

Etymology. This species is named for Bryant 
Mather in recognition of his long dedication to collect- 
ing Lepidoptera. 

Biology. Larval foodplants are unknown. March— 
November adult capture dates in the southern part of 
range suggest a bivoltine life cycle there; May and 
June capture dates in the northern part of range sug- 
gest univoltinism there. In the north, this species flies 
earlier than A. angulatana. 


Aethes terriae Sabourin & Miller, 
new species 
(Fig. 9a—c) 


Male exterior. Head, tegulum, and notum white, 
labial palpus 2x longer than horizontal eye diameter, 
second segment white apically. Fore- and midlegs 
black anteriorly with buff scales at junctions of tarsi, 
the longer tibial spur of middle leg buff; hindleg pre- 
dominately buff, a few dark scales on the shorter tibial 
spur. FW (Fig. 9a) 5.3 mm long (n = 3). Ground color 
white; markings black or buff with scattered buff 
scales; minute striae on costal and anal margins; costa 
black to basal fascia; basal fascia extending to costa, or 
obscure just below costa; IM fascia short, not extend- 
ing into discal cell, a few buff scales on apical margin; 
anal crescent narrow, buff with black borders; discal 
spot obscure; OM fascia present; ST line buff with 
black border, extending only to vein M,; fringe white 
with scattered buff and brown scales. HW pale drab; 
fringe paler; basal band present. 

Male genitalia (Fig. 9b, c) (n = 3). Costa of valva 
sinuate, posterior margin rounded, bearing a hooklike 
process ventrally; sacculus rounded. Median process 
of transtilla short, triangular. Sicklelike apical struc- 
tures of socii short. Aedeagus apically falcate, vesica 
bearing one long cornutus. Anellus lobes absent. Juxta 
a small subrectangular plate. 

Types. Holotype (Fig. 9a): MICHIGAN, Kalama- 
zoo Co., wet deciduous forest east side Sugarloaf Lake, 
3, 07/03/1993 (G. J. Balogh), FW 5.0 mm long, gen. 
prep. MS 99200 (AMNH). Paratypes (n = 2): 
MICHIGAN: Cass Co., Westrate farm, LT, 4, 
07/16/1999 (J. Vargo), gen. prep. MSO01088 (Fig. 9b, 
c), wing slide MSO1088W (UMSP): d, same data as 
holotype, gen. prep. MSO1069 (GJB). 


Diagnosis. Superficially, A. terriae resembles a small 
A. sexdentata. It differs from A. sexdentata superficially 
in its IM fascia not extending beyond discal cell, nar- 
rower anal crescent, and ST line not reaching apex (Figs. 
Ya, 5a). In male genitalia, A. terriae differs from A. sex- 
dentata in that the posterior margin of its valva is evenly 
rounded, with a hooklike spine on the ventral margin, 
transtilla with a short median process, an apically falcate 
aedeagus, and a single cornutus (Fig. 9b, c); in contrast, 
the valva of A. sexdentata has a more narrowly rounded 
posterior margin with a few spines near the outer margin 
of the harpe, a larger median process of transtilla, a small 
bundle of cornuti in addition to one large cornutus, and 
the aedeagus lacks a falcate apex (Fig. 5b, c). 

Discussion. The female of A. terriae is unknown. 

Etymology. This species is named in honor of Terri 
Balogh for her hospitality to lepidopterists. 

Biology. Larval foodplants are unknown. July 3-16 
adult capture dates suggest a univoltine life cycle. 


Aethes baloghi Sabourin & Metzler, 
new species 
(Fig. 10a-d) 


Male and female exterior. Head and notum 
mixed cream and buff; labial palpus 2x longer than 
horizontal eye diameter, apex cream colored. FW (Fig. 
10a) 4.0-6.8 mm long (n = 155). Ground cream suf- 
fused with buff; markings orange brown; fasciae dark 
bordered except for basal fascia; basal fascia oblique, 
costal half indiscernible; basal 1/4 of costa same color 
as markings; costal and anal margins with minute 
striae; IM fascia extending through cell, joined with 
discal spot, angled toward tornus; OM fascia reduced 
to a few scales near costa; ST line not conspicuous, 
consisting of a few scattered dark scales not reaching 
apex; fringe of ground color, but orange-brown on tor- 
nus adjacent to anal crescent. HW dark drab; fringe 
cream with a darker basal line. 

Male genitalia (Fig. 10b, c) (n = 15). Costa of valva 
fairly straight, posterior margin digitiform; sacculus 
rounded. Median process of transtilla long, broad, ta- 
pering terminally to a twisted apex. Aedeagus apically 
faleate, vesica bearing a scobinate patch, but no cor- 
nuti. Anellus lobes lateral and asymmetrical, inner mar- 
gin spinose terminally. Juxta broad, crescent shaped. 

Female genitalia (Fig. 10d) (n = 14). Anterior and 
posterior apophyses subequal in length to papillae 
anales. Sterigma, antrum, and ductus bursae reduced; 
sterigma a C-shaped sclerotized ridge; corpus bursae 
large, bulbous. Signum a sclerotized area on caudal 
right half of corpus bursae when viewed as in Fig. 10d. 
Surface of sternum 7 with heavily sculpted, irregular 
ridges. Cervix bursae indiscernible. 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Types. Holotype (Fig. 10a): NEW YORK, Yonkers 
[Westchester Co.], 2, 06/19/1936 (A. B. Klots), FW 6.5 
mm long, gen. prep. MSO0061 (AMNH). Paratypes 
(n = 154): ILLINOIS: Waukegan [Lake Co.], 2 
08/17/1941 (A. K. Wyatt), 309 (FMNH). Putnam Co., 
3, 09/20/1959: 3, 09/17/1964 (M. O. Glenn) (USNM). 
INDIANA: Lagrange Co., LT, 3, 07/26/1996. St. 
Joseph Co., LT, °, 08/07/1999, gen. prep. 00424IG, 
wing slide MS00424W (UMSP): 3 6, 07/28/1999: 3 4, 
08/26/1999: 3, 08/26/1999, gen. prep. EHM253; 3 3, 
08/28/1999 (J. Vargo) (JV). Hessville [Lake Co.], 2 
06/30/1905 (A. Kwiat), gen. prep. IG (USNM). MAS- 
SACHUSETTS: Martha’s Vineyard [Dukes Co.], d, 
08/11/1944 (F. M. Jones), gen. prep. IG (USNM). 
MICHIGAN: Allegan Co., T2N R14W Sec. 7, 2 d, 
05/22-24/1970 (J. P. Donahue), gen. prep. 
MS97345IG (MSUC): T2N R15W Sec. 25, d, Allle- 
gan] S[tate] G[ame] A[rea], 07/21/1984; T3N RI5W 
Sec. 36, °, 05/03/1991, gen. prep. IG; T2N R14W Sec. 
26, 2°, 09/09/1989; 4 6, 2 2 07/25/1987: 6, 2 
09/19/1989, gen. prep. EHM251: sand prairie, sa- 
vanna, 56 6, 15 2, 08/22/1986, gen. preps. IG, 
MS97350, EHM250, 252; 2 3, 09/19/1987, gen. prep. 
MS97347; T3N R14W Sec. 31, d, 05/11/1991; sand 
prairie/savanna, 9, 2 4, ey: gen. prep. 
EHM254 (G. Balogh) cae A{llegan] C[ounty] 
G[ame] P[reserve], LT, 2 4, 07/17/1999, gen. prep. 
MS99322; Alllegan] ae G[ame] Alrea], 8 4d, 
05/26/2000 (J. Vargo), gen. preps. 004231G, MS01083 
(Fig. 10b) (UMSP). Cheboygan Co., Grass Bay, T38N 
RSW Sec. 25, 6, 08/04/1995, gen. prep. IG (BGS); 
T37N R38W Sec. 33, 2, 06/25/1990 (B. G. Scholtens), 
gen. prep. MS99154 (UMSP). Emmet Co., Wilderness 
S[tate] Plark], base of Waugoshance Point, 4, 
05/27/1990, gen. prep. IG (G. Balogh) (GJB). Lk 
Lansing [Ingham Co.], 3 2, 09/20/1936, gen. prep. 
MS01097; 3 d, same data (W. C. Stinson), gen. preps. 
IG (MSUC). Mackinaw Co., T41N R5W Sec. 23, 2, 
08/15/1998, dunes (G. Balogh) (GJB). Montcalm Co., 
T12N R10W Sec. 18, 4, 05/09/1987 (G. Balogh), gen. 
prep. IG. Muskegon Co., T12N R17W Sec. 26, sand 
priairie)/savanna, 3 ¢, 06/11/1993, gen. prep. IG; 
Muskegon S[tate] Plark] dunes, 3, 07/13/1991; 4, 
08/12/1989: 3d, 07/15/1991 (G. Balogh) (GJB). 
Newaygo Co., T12N R12W Sec. 1, d, 07/14/1967 (J. P. 
Donahue), gen. prep. MS01098 (Fig. 10c). Wayne Co., 
°, 09/14/1943 (J. H. Newman), gen. prep. IG (MSUC). 
MISSISSIPPI: Harrison Co., Long Beach, 9, 03/07/ 
1991, gen. prep. IG, coll. #180535: d, 03/20/1998, gen. 
prep. IG, coll. #180537 (BMMS); 6, 03/13/1991, gen. 
prep. IG, coll. #180534; 2, 04/23/1991, gen. prep. IG, 
coll. #180553; d, 04/30/1997, gen. prep. IG, coll. 
#180548 (R. Kergosien) (UMSP). MISSOURI: Barry 


VOLUME 56, NUMBER 4 


Cou Roaming River S[tate] Plark], Ozark ee 
flora, BL, °, 09/03/1994, gen. prep. MS99178 (RL); ° 
same data, gen. prep. MS99180 (UMSP). @olamnbia 
[Boone Co.], °, 09/17/1983 (W. S. Craig), gen. prep. 
MS98164 (UM). NEW JERSEY: Franklinville 
[Gloucester Co.], 2, 08/18/1939 (W. Sachse). Lake- 
hurst [Ocean Co.], 9°, 09/04/1937, gen. prep. 
MS01054; AL, d, 08/08/1939. New Lisbon [Burling- 
ton Co.], AL, 4, 09/23/1935; 2, 07/06/1942, gen. prep. 
MS00240; Whitesbog, AL, 2, 07/31/1938 (E. P. Dar- 
lington), gen. prep. MSO1077 (Fig. 10d)(ANSP). 
NEW YORK: Yonkers [Westchester Co.], 4, 
06/06/1936; 2, 09/14/1936 (Klots) (AMNH). NORTH 
CAROLINA: So[uthern] Pines [Moore Co.], 9, 
04/1-7, gen. prep. IG. VIRGINIA: Glencarlyn [Ar- 
lington Co.], 2, AL, 05/26/1959 (J. Armstrong), gen. 
prep. IG (USNM). 

Diagnosis. A. baloghi most resembles A. patricia 
Metzler, but differs in having orange-brown FW mark- 
ings compared to the buff markings of A. patricia. In 
male genitalia of A. baloghi the posterior margin of the 
valva is digitiform, not outlined with dense setae, the 
sacculus is undemarcated, and anellus lobes are pres- 
ent (Fig. 10b, c); in contrast, A. patricia has the poste- 
rior margin of the valva concave, outlined with dense 
setae, tipped with a short tooth, the sacculus as a sepa- 
rate lobe, and anellus lobes are absent (Metzler 2000: 
fig. 2a, b). In female genitalia, A. baloghi has a con- 
spicuous signum and sculptured aan sternum (Fig. 
10d), whereas A. patricia lacks a signum, and sternum 
7 has two heavily sclerotized areas laterad to the os- 
tium bursae (Metzler 2000: fig. 2c). 

Discussion. Males were associated by FW color 
pattern and simultaneous capture. 

Etymology. This species is named for George J. 
Balogh in recognition of his dedication to lepi- 
dopterology. 

Biology. Larval foodplants unknown. Associated 
with prairie habitat in parts of range. March—Septem- 
ber adult capture dates suggest a multivoltine life cycle. 


Aethes atomosana (Busck) 
(Fig. 1la-d) 


Phalonia atomosana Busck (1907:22) (Holotype ° [in- 
correctly given as ¢ by Brown & Lewis 2000], type 
#10223, Pa., Pittsburgh [Allegheny Co.], H. Engel, 
08/17/1905, forering 8.0 mm long, gen. prep. 
USNM23825, in USNM), Forbes (1923:509), Mc- 
Dunnough (1939:59), Procter (1946:308), Brower 
(1983:49, in part), Handfield (1997:44), Brown and 
Lewis (2000:1020). 

Aethes atomosana; Grehan et al. (1995:25), Razowski 
(1997:127, figs. 82-84, 163-165). 


231 


Phtheochroa atomosana; Poole and Gentili (1996:876), 

Nielsen (1998:10). 

Male and female exterior. Head, notum, and 
tegulum chamois colored; labial palpus 3.0—3.5x 
longer than horizontal eye diameter. Thoracic tuft 
conspicuous. FW (Fig. lla) 6.6—9.8 mm long (n = 
34). Ground color chamois; markings reduced, rep- 
resented by black and buff scales; some specimens 
completely irrorated with fuscous scales, others less 
maculate with no markings above vein Sc; fuscous 
striae along anal margin and distally on costa beyond 
vein Sc; basal fascia vestigial, serial black spots along 
vein Sc; IM and OM fasciae absent; discal spot pres- 
ent; anal crescent reduced to a submarginal spot; ST 
line represented by paired spots reaching apex; 
fringe of ground color; heavy fuscous scaling on un- 
derside excluding margins. HW cream to fawn col- 
ored. 

Male genitalia (Fig. 11b, c) (n = 10). Costa of valva 
slightly curved, posterior margin subsinuate, oblique, 
serrated, with apical spine; sacculus rounded, forming 
a subharpal plate. Median process of transtilla large, 
thumb shaped, apex broadly rounded, with 4 or more 
short apical processess. Aedeagus bearing a medial 
process, vesica bearing one long, thin cornutus. Anel- 
lus lobes bilateral and appressed. Juxta large, subovate. 

Female genitalia (Fig. 11d) (n = 4). Tete post- 
vaginalis a long elliptical plate; anterior apophyses 
longer than posterior; stemum 7 heavily sclerotized, a 
pair of raised ridges creating a medial furrow ventrad 
of antrum; antrum cylindrical, % as long as ductus bur- 
sae. Cervix bursae indiscernible. 

Diagnosis. A. atomosana most resembles A. 
matthewcruzi, which is described and differentiated in 
the next section. 

Biology. Larval foodplants unknown. Early August 
to mid-September adult captures suggest a univoltine 
life cycle. 

Specimens examined (n = 34, including type). 
Counties and months of capture by State or Province: 
Canada. NOVA SCOTIA: Kings, August; Lunenburg, 
August (MS, NSPM, UMSP). ONTARIO: Carleton, 
August (CNC). USA. ILLINOIS: Algonquin, August 
(INHS). INDIANA: Lagrange, August (JV). MARY- 
LAND: Garrett, September (JDG). MICHIGAN: 
Barry, September; Cass, September; Cheboygan, Au- 
gust; Clinton, September; Kalamazoo, August; Liy- 
ingston, September; Midland, August (BGS, GJB, 
MSUC, UMSP, WPW). PENNSYLVANIA: Allegheny, 
August (USNM). VERMONT: Addison, August; Chit- 
tenden, August, September; Franklin, August; Grand 
Isle, September (FMPS, MS, MSG). WISCONSIN: 
Burnett, August (MS). 


Aethes matthewcruzi Sabourin & Vargo, 
new species 


(Fig. 12a—d) 


Male and female exterior. Head cream with 
brown suffusion; labial palpus 2.5-3.0x longer than 
horizontal eye diameter. Notum and tegulum cream 
with brown suffusion or mixed black, buff, and brown; 
hindleg anterior spurs and shorter tibial spurs with 
some darker scales. FW (Fig. 12a) 6.9-9.2 mm long (n 
= 12). Ground color cream with brown suffusion espe- 
cially between veins; markings reduced to small, black, 
buff, and brown spots; basal “4 of costa gray, costa and 
anal margin with fuscous striae; IM and OM fasciae 
and anal crescent reduced to submarginal black spots; 
discal spot the most conspicuous marking, a mixture of 
black and buff scales; gray patch between discal spot 
and anal crescent extending to tornus; ST line repre- 
sented by black and buff spots, not extending to apex; 
fringe cream colored with basal line the color of FW 
markings; heavy fuscous scaling on entire underside. 
HW drab-gray. 

Male genitalia (Fig. 12b, c) (n = 5). Valva broad, 
costa slightly curved, posteriorly deeply emarginate, 
serrated; sacculus long, slightly curved, tapering dis- 
tally. Median process of transtilla triangular, apical half 
narrowing to a point. Phallobase narrow and curving to 
a 45° angle to aedeagus. Aedeagus thick, vesica bear- 
ing one long, thin cornutus and an elongate scobinate 
patch. Anellus lobes (Fig. 12c) long and narrow. 

Female genitalia (Fig. 12d) (n = 4). Sterigma sub- 
quadrate; lamella postvaginalis a medial proccess; 
apophyses long, anterior longer than posterior; antrum 
a robust collar, % as long as ductus bursae; ductus bur- 
sae sclerotized entire length. 

Types. Holotype (Fig. 12a): MINNESOTA, 
Roseau Co., Lost River S[tate] Florest], AL, ¢, 
08/22/1999 (Vargo & Sabourin), forewing length 8.0 
mm, gen. prep. MS00167 (Fig. 12b, c) (UMSP). 
Paratypes (n = 11): MAINE: Steuben [Washington 
Co.], 3, 08/02/1990 (M. A. Roberts), gen. prep. G1786 
(MAR). MICHIGAN: Allegan Co., Saugatuck S[tate] 
P[ark], 2, 08/08/1992, gen. prep. MS97269. Barry Co., 
fen on Shaw Lk., T3N R14W Sec. 3, 4, 06/26/1991 (G. 
Balogh), gen. ep MS01055 (GJB). Cass Co., T5S 
R14W Sec. 31, 2, 08/22/1996 (W. P. Westrate), gen. 
prep. MS01030 (WPW). Washtenaw Co., T28 R3E 
Sec. 6, 6, 07/26/1993 (J. H. Wilterding), gen. prep. 
MS00298, wing slide MS00298W (JHW). MIN- 
NESOTA: Same data as holotype, 4 4, gen. prep. 
MS99543 (MS). NEW HAMPSHIRE: Whitefield 
[Coos Co.], 2, 08/07/1985 (W. J. Kiel), gen. prep. 
MS97209 (UMSP). WISCONSIN: Burnett Co., 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Grantsburg, AL, 2, 07/28/1999, gen. prep. MS00342 
(Fig. 12d), wing slide MS00342W; T40N R1SW Sec. 
93, LT, 6, 07/14/2001 (M. Sabourin) (MS). 

Diagnosis. A. matthewcruzi is most similiar to A. 
atomosana, from which it differs subtly in FW mark- 
ings, and more markedly in genitalia. A. mattheweruzi 
has fuscous striae on the basal “4 of the FW costal 
margin which are absent in A. atomosana, and the OM 
and IM fasciae, which are reduced to submarginal 
spots in A. matthewcruzi, are lacking completely in A. 
atomosana (Figs. 12a, 1la). In male genitalia of A. 
matthewcruzi, the posterior margin of the valva is 
deeply emarginate, the sacculus long and slightly 
curved, and the median process of transtilla triangular, 
whereas in A. atomosana the posterior margin of the 
valva is acute, the sacculus short and rounded, and the 
median process of transtilla thumb shaped (Figs. 12b, 
11b). In female genitalia, A. matthewcruzi has a small, 
round lamella postvaginalis and sternum 7 is not heav- 
ily sclerotized, whereas in A. atomosana the lamella 
postvaginalis is elliptical, and sternum 7 is heavily 
sclerotized (Figs. 12d, 11d). Sexual dimorphism in size 
may occur, as male FW length ranged 6.8-8.2 mm (n = 
8), and female, 8.6-9.2 mm (n = 4). However, this 
might prove to be an artifact of small sample size. 

Discussion. Females were associated by FW color 
pattern. 

Biology. Larval foodplants unknown. June-August 
adult capture dates suggest a univoltine life cycle. 
Adults appear about one month earlier than those of 
A. atomosana. Although broadly sympatric with A. ato- 
mosana, A. matthewcruzi seems to occur in a wider va- 
riety of habitats. 

Etymology. This species is named in memory of 
Matthew Edward Cruz, a talented young artist who 
lost his life in a tragic accident. 

In conclusion, using male structural characters, we 
readily constructed a key to the 12 species treated, but 
it is omitted here because numerous untreated species 
of Aethes also occur in the region. 


ACKNOWLEDGEMENTS 


We thank the following for specimen loans or use of facilities: V. 
Albu, D. Azuma, G. J. Balogh, C. B. Barr, J. W. Brown, P. J. Clausen, 
W. S. Craig, P. T. Dang, P. Z. Goldstein, J. R. Grehan, M. S. Griggs, 
T. Harrison, J. R. Sore, W. Kiel, S. Krauth, R. D. Kriegel, B. 
Landry, J.-F. Landry, R. Letsinger, J. Lewis, B. Mather, K. Methven, 
M. C. Nielsen, M. F. OiBaen, J. A. Powell, R. J. Pupedis, E. L. 
Quinter, J. E. Rawlins, F. H. Rindge, M. A. Roberts, B. G. Scholtens, 
K. B. Simpson, R. W. Sites, F. W. ‘Gtehr. K. R. Tuck, W. P. Westrate, 
J. H. Wilterding, and B. Wright. We thank M. G. Pogue for making 
photographic slides of type specimens available, R. W. Holzenthal 
for use of photomicrographic equipment, B. J. Sabourin for photo- 
graphic assistance, C. E. Weyland for logistical assistance, and J. A. 
Powell, G. J. Balogh, M. A. Roberts, M. G. Pogue, and J. W. Brown 
for useful manuscript reviews. 


VOLUME 56, NUMBER 4 


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Received for publication 4 December 2001; revised and accepted 26 
May 2002. 


Journal of the Lepidopterists’ Society 
56(4), 2002, 234-264 


SYSTEMATICS OF MOTHS IN THE GENUS CATOCALA (NOCTUIDAE). III. THE TYPES OF 
WILLIAM H. EDWARDS, AUGUSTUS R. GROTE, AND ACHILLE GUENEE 


LAWRENCE F. GALL 


Entomology Division, Peabody Museum of Natural History, Yale University, New Haven, Connecticut 06511, USA 


AND 


Davip C. Hawks 


Department of Entomology, University of California, Riverside, California 92521, USA 


ABSTRACT. Nomenclatural status is assessed for the 75 Catocala names authored by William Henry Edwards, Augustus Radcliffe Grote 
(including Coleman T. Robinson), and Achille Guenée. Three neotypes s and 46 lectotypes are designated, and six new or reniced synonymies are 
presented. Historical and biographical notes on these authors and the collections on w yhich they ‘based their descriptions are provided, Addi- 
tionally, the unpublished Crise ala paintings by the early North American naturalist John Abbot are analyzed in detail for the first time. 


Additional key words: 


The holarctic genus Catocala Schrank (1802) is one 
of the most species-rich genera in the large moth 
family Noctuidae, with over 210 species split approxi- 
mately equally between the Nearctic and Palearctic re- 
gions. Despite the fact that Catocala are large, color- 
ful, and common moths that have been popular with 
lepidopterists for centuries, there has been no defini- 
tive systematic revision published for the entire genus, 
and the most current work treating all the Nearctic 
species is now approaching nearly a century old (Barnes 
& McDunnough 191Sb; the most recent checklist cov- 
ering the Nearctic taxa is Hodges et al. 1983). 

The present paper is the last in a series on the tax- 
onomy of the Nearctic Catocala appearing in advance 
of a Fascicle on the genus for the Moths of America 
North of Mexico. In the first two papers in the series 


(Gall 1990, Gall & Hawks 1990) we addressed 56 of 


the 347 names comprising the synonymy of the Nearc- 
tic Catocala, focusing on type material at the Field 
Museum of Natural History (Chicago) and the Mu- 
seum of Comparative Zoology (Harvard University), 
and designating 32 lectotypes. Herein, we address 75 
of the remaining Nearctic Catocala names authored by 
William H. Edwards, Augustus R. Grote, and Achille 
Guenée during the 19th century, designating 46 lecto- 
types and 3 neotypes, and establishing six new or re- 
vised synonymies and status amendments. Table 1 
summarizes our taxonomic decisions. 

This paper is divided into separate sections for Ed- 
wards, Grote, and Guenée, to highlight the biographi- 
cal and type disposition issues particular to each au- 

thor. Each section treats available names (species and 
subspecies) and then unavailable names s (infrasubspe- 
cific entities such as “aberration’”), respectively, in al- 
phabetical order. Although names deemed to be infra- 
imavailable under the International 
Code of Zoological Nomenclature (ICZN 2000), and 


subspecifie are 1 


taxonomy, neotypes, lectotypes, collections, history, biography. 


do not require formal treatment, we treat them here 
because we feel omitting them is a false economy that 
ultimately hampers revisory work, especially for 
groups like Catocala with lengthy and complex syn- 
onymies and historical literatures. 


NOMENCLATURAL PROTOCOL 


The terms “form” and “variety” were used habitually 
in descriptions of new taxa by ‘the Nearctic C atocala 
workers of the late 19th and early 20th centuries, and 
the tradition in the genus at that time was to apply 
these two terms in an infrasubspecific manner. How- 
ever, without knowing this custom, it is often impossi- 
ble to deduce infrasubspecific intent from the texts of 
the original descriptions alone, and the custom was cer- 
tainly not applied universally. Fortunately, the earlier 
Catocala monograph by Barnes and McDunnough 
(1918b) and the ‘lapidannten an checklists of Barnes and 
McDunnough (1917) and McDunnough (1938) indi- 
cate the availability previously deduced for such names. 
Because this complex and species-rich genus has never 
been fully revised, we opted to reassess availability for 
each “form” and “variety” name. The reason we did this 
is that infrasubspecific names, which are otherwise ex- 
cluded from zoological nomenclature, become formally 
available if they are used later in a clearly subspecific or 
specific manner. In such instances, the name takes the 
date and authorship of the person(s) who used it at the 
elevated rank (see Articles 10.2 and 45.6 of the Code). 
Thus, if an original description of a Catocala suggested 
subspecific rank according to Article 45.6.4, but the 
Barnes and McDunnough monograph and checklists 
indicated that infrasubspecific rank had been previ- 
ously deduced, we traced the name through the litera- 
ture to insure that infrasubspecific usage had indeed 
been intended and had remained consistent. Discrep- 
ancies are treated in the accounts for individual names. 


VOLUME 56, NUMBER 4. 


The terms “type” and “types” were also used in the 
majority of the original descriptions of Nearctic Cato- 
cala, often loosely. Because the texts for many of the 
descriptions could be termed telegraphic at best, we 
assumed the existence of syntypes when the singular 
“type” was used but the number of specimens was not 
otherwise readily and explicitly determinable. Often, 
type specimens for particular names were present in 
two or more institutional collections. We usually found 
evidence leading us to favor material from one institu- 
tion, either on the basis of the descriptions per se, 
specimen labels, or related published information. The 
order of preference among institutional collections 
varies by author, and we discuss these and related idio- 
syncracies (eam missexed specimens) as appropriate. 
When we were unable to determine a precedence 
among available syntypes, we normally selected the 
specimen from the institution containing most of the 
author's collection. 

In most species of Nearctic Catocala there is com- 
plex individual variation in wing pattern, as well as par- 
allel polymorphs that occur in both closely and more 
distantly related species. Largely because of this, a 
number of species boundaries among Nearctic Cato- 
cala remained poorly understood until our recent field 
and rearing studies covering the entire fauna. More- 
over, many Catocala species are simply difficult to dis- 
tinguish from one another, even when species’ bound- 
aries are well known (indeed, a number of the syntypic 
series for names in this genus contain more than one 
Catocala species). These problems are especially man- 
ifest in the group of large, pink-hindwinged Catocala 
that feed as larvae on willows and poplars. The princi- 
pal issue with the willow/poplar feeding taxa is that the 
extent of infrapopulational variation is far greater than 
previously understood. The group in fact consists of a 
much smaller array of species than indicated in the 
most recent faunal checklist by Hodges et al. (1983). A 
large number of the available names is referable to 
only five wide-ranging, morphologically variable, and 
difficult to separate Rocky Mountain and Pacific coast 
species: faustina Strecker, hermia Hy. Edwards, cali- 
fornica Edwards, semirelicta Grote, and junctura 
Walker. Because these willow/poplar feeders also tend 
to have the most tangled nomenclatural histories, we 
feel it is imperative to fix all the involved names firmly. 
Thus, in this paper, we have designated a neotype for 
any available name that refers to a willow/poplar feed- 
ing taxon for which original type material is apparently 
no longer extant. 

Nearctic Catocala type specimens are distributed in 
many institutional collections. However, the over- 
whelming majority of these types are at only eight in- 


stitutions: the Academy of Natural Sciences of 
Philadelphia (ANSP), the American Museum of Nat- 
ural History (AMNH), the Carnegie Museum of Nat- 
ural History (CMNH), the Field Museum of Natural 
History (FMNH), the Museum of Comparative Zool- 
ogy (MCZ), the Natural History Museum (London) 
(BMNH), the National Museum of Natural History 
(USNM), and the Yale Peabody Museum of Natural 
History (YPM). Types of Catocala are maintained in 
separate lepidopteran type collections at most of these 
institutions, but remain integrated with the general 
collections at others (e.g., the Strecker material at the 
FMNH, and the main historical series at the BMNH). 


WILLIAM HENRY EDWARDS 


W. H. Edwards was one of the most highly ac- 
claimed American lepidopterists of the 19th century. 
His life and accomplishments have been well chroni- 
cled (see e.g., Mallis 1971), and he is probably best 
known for his monumental three-volume treatise The 
Butterflies of North America. W. H. Edwards was also 
the first American to focus on the Nearctic Catocala, 
naming a total of 10 new species in 1864. 

During the 19th century, many of Edwards’ Cato- 
cala types apparently were extant in Philadelphia in 
the American Entomologial Society collection, but by 
the time Beutenmiiller was working on his revision of 
the genus, many of the types had vanished. Beuten- 
miiller (1918a:44) stated: “These were supposed to be 
with the American Entomological Society, but I could 
not find them there nor in any other collection. I con- 
sequently wrote to W. H. Edwards shortly before his 
death [in 1909] asking for information on the matter 
and he informed me that all his Catocala went to Mr. 
Julius Meyer, Brooklyn, N.Y. After Mr. Meyer's death 
his collection was bought by the Kny-Scheerer Co., 
New York, and Dr. G. Lagai writes me that the Cato- 
cala part of the same was sold to Hon. Walter Roth- 
schild, Tring, London, England. If Edwards’ types are 
still extant the same will probably be found in the 
Rothschild collection.” 

We have searched throughout the North American 
catocalines at the BMNH, and have not found any def- 
inite Edwards types in either the main systematic, 
Rothschild, or Oberthur collections. However, in the 
“Mixed Authors” type area at the BMNH is a drawer 
containing 34 specimens of 16 Catocala species, in- 
cluding representatives of each of the Catocala au- 
thored by Edwards, except for similis Edwards. The 
material in this drawer had been segregated out of the 
Rothschild collection. A single typed sheet accompa- 
nies this drawer, on which are listed Edwards’ and 
Meyer's Catocala names, along with the statement: 


236 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Synopsis of taxonomic assessments and actions taken in this paper pertaining to names in the moth genus Catocala Schrank. Insti- 
tutional abbreviations are as listed in the text section Nomenclatural Protocol; additionally, RUPM = Roemer und Pelizaeus Museum, Germany, 
NYSM = New York State Museum, Albany. Note under the holotypes that a petition (Case 3210) is currently pending at the ICZN to suppress 
polygama Guenée, an unused senior synonym of alabamae Grote. 


Taxon Author Date Location Rank assigned herein Comments on rank 
PREVIOUSLY DESIGNATED HOLOTYPES 

arizonae Grote 1873 ANSP synonym of junctwra Walker (1858) new synonymy 
beaniana Grote 1878 BMNH synonym of meskei Grote (1873) no change 
cara Guenée 1852 USNM full species no change 
clintoni Grote 1864 CMNH full species no change 
coelebs Grote 1874 BMNH subspecies of badia Grote & Robinson (1866) new status 
editha Edwards W. H. 1874 ANSP synonym of amatrix (Hiibner 1813) new synonymy 
innubens Guenée 1852 USNM full species no change 
insolabilis Guenée 1852 USNM? full species no change 
lacrymosa Guenée 1852 USNM full species no change 
marmorata Edwards W. H. 1864 ANSP? full species no change 
messalina Guenée 1852 USNM? full species no change 
moderna Grote 1900 RUPM synonym of maestosa Hulst (1884) no change 
muliercula Guenée 1852 USNM? full species no change 
polygama Guenée 1852 USNMP? synonym of grynea (Cramer 1780) no change 
uxor Guenée 1852 USNM? synonym of ilia (Cramer 1780) no change 


PREVIOUSLY DESIGNATED NEOTYPES 


californica Edwards W. H. 1864 CMNH full species no change 
PREVIOUSLY DESIGNATED LECTOTYPES 

adoptiva Grote 1874 MCZ synonym of delilah Strecker (1874) no change 
angusi Grote 1876 AMNH full species no change 


NEOTYPES DESIGNATED HEREIN 


connubialis Guenée 1852 BMNH full species no change 
micronympha Guenée 1852 AMNH full species no change 
walshii Edwards 1864 ANSP synonym of junctura Walker (1858) no change 
LECTOTYPES DESIGNATED HEREIN 

abbreviatella Grote 1872 ANSP full species no change 
alabamae Grote 1875 BMNH full species no change 
andromedae Guenée 1852 illustration full species no change 
androphila Guenée 1852 USNM synonym of amica (Hiibner 1813) no change 
anna Grote 1874 BMNH synonym of amestris Strecker (1874) no change 
badia Grote & Robinson 1866 BMNH full species no change 
briseis Edwards W. H. 1864 ANSP full species no change 
cerogama Guenée 1852 USNM full species no change 
chelidonia Grote 1881 USNM full species no change 
coccinata Grote 1872 ANSP full species no change 
communis Grote 1872 ZMHU synonym of neogama (J. E. Smith 1797) no change 
desperata Guenée 1852 USNM synonym of vidua (J. E. Smith 1797) no change 
dulciola Grote 1881 AMNH full species no change 
flebilis Grote 1872 ANSP full species no change 
formula Grote & Robinson 1866 ANSP synonym of similis Edwards (1864) no change 
fratercula Grote & Robinson 1866 ANSP synonym of micronympha Guenée (1852) revised synonymy 
frederici Grote 1872  ZMHU full species no change 
gracilis Edwards W. H. 1864 YPM full species no change 
habilis Grote 1872 ANSP full species no change 
levettei Grote 1874 BMNH synonym of judith Strecker (1874) no change 
lineella Grote 1872 BMNH full species no change 
melanympha Guenée 1852 USNM synonym of antinympha (Hiibner 1823) no change 
meskei Grote 1873 USNM full species no change 
minuta Edwards W. H. 1864 YPM full species no change 
mira Grote 1876 BMNH full species no change 
palaeogama Guenée 1852 USNM full species no change 
parta Guenée 1852 USNM full species no change 
parvula Edwards W. H. 1864 ANSP synonym of minuta Edwards (1864) no change 
phalanga Grote 1864. BMNH synonym of palaeogama Guenée (1852) no change 


piatrix Grote 1864 CMNH full species no change 


VOLUME 56, NUMBER 4 


Taxon 


Author 


Date 


Location 


LECTOTYPES DESIGNATED HEREIN (Continued) 


TABLE I. 


Rank assigned herein 


bo 
(os) 
~l 


Continued. 


Comments on rank 


ponderosa Grote & Robinson 1866 BMNH synonym of nebulosa Edwards (1864) no change 

praeclara Grote & Robinson 1866 CMNH full species no change 

residua Grote 1874 AMNH full species no change 

retecta Grote 1872 ANSP full species no change 

robinsoni Grote 1872 ANSP full species no change 

scintillans Grote & Robinson 1866 BMNH synonym of innubens Guenée (1852) no change 

semirelicta Grote 1874 BMNH full species no change 

similis Edwards W.H. 1864 YPM full species no change 

simulatilis Grote 1874 BMNH synonym of obscura Strecker (1873) no change 

sinuosa Grote 1879 BMNH subspecies of coccinata Grote (1872) no change 

snowiana Grote 1876 AMNH synonym of palaeogama Guenée (1852) revised synonymy 
sordida Grote 1877 BMNH full species no change 

subnata Grote 1864 BMNH full species no change 

tristis Edwards W. H. 1864 ANSP synonym of andromedae Guenée (1852) revised synonymy 
verrilliana Grote 1875 BMNH full species no change 

westcottii Grote 1878 BMNH synonym of amestris Strecker (1874) no change 

NO ACTION 

basalis Grote 1876 BMNH synonym of habilis Grote (1872) infrasubspecific name 
bunkeri Grote 1876 BMNH synonym of cerogama Guenée (1852) infrasubspecific name 
flavidalis Grote 1874 BMNH synonym of innubens Guenée (1852) infrasubspecific name 
guenei Grote 1887 no type replacement name for viduata Guenée (1852) no change 

hilli Grote 1883 NYSM synonym of concumbens Walker (1858) infrasubspecific name 
nebulosa Edwards W. H. 1864 ANSP? full species no change 

serena Edwards W. H. 1864 ANSP? full species no change 

viduata Guenée 1852 no type replacement name for vidua (J. E. Smith 1797) no change 


“The following are W. H. Edwards’ species which 
Beutenmiiller stated went to the Meyer Coll. and 
types ought to be in Tring.” The typed sheet is un- 
dated, but almost certainly predates 1974 (M. Honey 
pers. com. 1999). 

A type of Catocala gisela Meyer is in this drawer, 
and this suggests the material is indeed from Meyer, 
and is that refered to by Beutenmiiller (1918a). The 
representative specimens of Edwards’ species in the 
drawer all bear labels in what appears to be A. Rogen- 
hofer’s handwriting, giving the collecting locality and 
other details, as well as stating “Type.” However, the 
label data on most of these supposed Edwards types 
do not correspond to what appears in the respective 
original descriptions by Edwards. For example, the 
specimen of Catocala marmorata Edwards is labelled 
“Ky.” (=Kentucky) whereas the original description 
calls for Yreka, California, and its abdomen is also in- 
tact, contradicting Strecker (1874:73), who figured the 
marmorata type and stated that the “abdomen is want- 
ing in the single example so far known;” the specimen 
of Catocala briseis Edwards is labelled “Canada” 
whereas the original description calls for one specimen 
from the Catskills [New York] and one specimen from 


Rhode Island, and both of these two syntypes have al- 
ready been located in other collections; and so forth. 
Thus, we feel these BMNH specimens are not Ed- 
wards’ types, but rather are specimens that were con- 
sidered to be typical, or compared to the types (or per- 
haps even asserted to be types during the several 
transfers of Edwards’ material, to enhance their mon- 
etary value). 

Although Beutenmiiller (1918a) indicated he could 
not find Edwards’ Catocala types at the ANSP, we 
readily located one of his two briseis syntypes in the 
main ANSP collection that apparently had not previ- 
ously been segregated into the type cabinets. We have 
also explored the CMNH collection for possible Ed- 
wards Catocala types, with limited success (see our 
notes on the ANSP and CMNH below under A. R. 
Grote). Another collection known to have been uti- 
lized heavily by Edwards, Grote, and Strecker in the 
1860's and 1870's was that of Mrs. Bridgham, from 
Rhode Island. In 1991, the senior author located the 
Bridgham family’s collection at the Smithfield branch of 
the Audubon Society of Rhode Island, and its specimen 
material was transferred to the Yale Peabody Museum 
of Natural History in 1992 (about 1800 specimens were 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fic. 1. ‘Type specimens of moths in the genus Catocala Schrank. W. H. Edwards types. a, lectotype, briseis. b, neotype, californica. e, lec- 
totype, gracilis. d, lectotype, minuta. e, lectotype, parvula. f, lectotype, similis. g, lectotype, tristis. h, neotype, walshii. 0.9-1.0 x natural size. 


salvaged, and the Victorian-era cabinet remained with 
the Audubon Society; Gall 1995). A number of the 
Bridgham specimens bear Grote determination labels, 
and syntypes for all Catocala species that Edwards 
(1864) cited explicitly as coming from the Bridgham 
collection were extant and identifiable. The Bridgham 
collection is almost unquestionably an uncited source 
for all Catocala that Edwards authored that are native 
to the eastern USA. Our procedure for W. H. Ed- 
wards’ names is therefore to give precedence first to 
type material from any explicitly cited collection, then 


to Bridgham (YPM), and then to ANSP/CMNH. 


Available Names 


Catocala briseis W. H. Edwards, 1864. Proc. Ento- 
mol. Soc. Phil. 2:508 


The original description states “I have seen but two 
individuals of this species, one of which was taken by 
me in the Catskills in July, 1861; the other is in the fine 
collection of Mrs. Bridgham, taken in Rhode Island.” 
One unlabeled male ex Bridgham collection is at the 
YPM, and a female labeled as briseis by Edwards is at 
the ANSP. We give precedence to the labeled ANSP 
female, and to clarify application of the name we 


VOLUME 56, NUMBER 4 


hereby designate it as LECTOTYPE for briseis (Fig. 
la). The lectotype bears the labels “briseis” and “LEC- 
TOTYPE/Catocala briseis/Edwards 1864/Desig. Gall 
& Hawks 2002.” The type locality is hereby restricted 
to the Catskill [Mountain ]s, [New York, USA]. We rec- 
ognize Catocala briseis as a full species. 


Catocala californica W. H. Edwards, 1864. Proc. En- 
tomol. Soc. Phil. 2:509 


The original description does not state the number 
of types. Although Strecker claimed to have figured 
the type of californica, Barnes and McDunnough 
(1918b:24-25) provided credible evidence that 
Strecker’s specimen was not from the type series. They 
also located a specimen at the CMNH with suitable lo- 
cality data “which proved to be what has generally 
been known as mariana |Hy.] Edwards,” and stated 
“We propose, therefore, to consider this specimen as 
typifying the true californica and list mariana as a syn- 
onym.” We consider Barnes and McDunnough’s dis- 
cussion and subsequent proposal to constitute a valid 
neotype designation for californica, meeting the crite- 
ria listed in Article 75 of the Code. We show their neo- 
type in Fig. lb. The type locality is Yreka, [Siskiyou 
County,] California [USA]. We recognize Catocala cal- 
ifornica as a full species. 


Catocala editha W. H. Edwards, 1874. Trans. Amer. 
Entomol. Soc. 5:112 


The original description states “a single specimen 
taken in Sonoto Valley, by Mr. Henshaw, July, 1874.” 
This specimen, the holotype by monotypy, is at the 
ANSP (ANSP Type #7719). The type locality is Sonoto 
Valley, [Arizona, USA]. The name editha has previ- 
ously been tabulated in the Nearctic literature as a 
subspecies of Catocala amatrix (Hiibner, [1813]). 
However, the editha holotype falls within the normal 
range of variation seen in nominate amatrix from else- 
where in North America (the infrasubspecific form 
“pallida” Barnes & McDunnough (1918b), with sil- 
very-white forewing shading and a reduced hindwing 
medial band, is usually listed as a synonym of editha; 
specimens of “pallida” occur sporadically in the west- 
ern Plains states and more consistently in the Rocky 
Mountains). We recognize Catocala editha as a syn- 
onym of Catocala amatrix (Hiibner) (NEW SYN- 
ONYMY). 


Catocala gracilis W. H. Edwards, 1864. Proc. Ento- 
mol. Soc. Phil. 2:511 


The original description does not state the number 
of types, but indicates “Mrs. Bridgham’s Collection.” A 
series of 12 gracilis ex Bridgham collection is at the 


239 


YPM. One male bears a handwritten label “gracilis? 
Edw.” in Grote’s handwriting, unquestionably dating 
from around the time of Grote’s stated uncertainty 
about the name (Grote 1877:169-170). To clarify ap- 
plication of the name we hereby designate this male as 
LECTOTYPE for gracilis (Fig. 1c). In addition to the 
above label, the lectotype bears the labels “Samuel 
Willard Bridgham/Collection/Presented Rhode Is- 
land/Audubon Society, May 1992” and “LECTO- 
TYPE/Catocala gracilis/Edwards 1864/Desig. Gall & 
Hawks 2002.” The type locality is not specified, but is 
probably Rhode Island, USA. We recognize Catocala 
gracilis as a full species. 


Catocala marmorata W. H. Edwards, 1864. Proc. En- 
tomol. Soc. Phil. 2:508 


Of the type of marmorata, Strecker (1874:73) rather 
amusingly stated “abdomen is wanting in the single ex- 
ample so far known . . . the unique type from which 
the annexed figure was drawn is in the Museum of the 
Am. Ent. Soc.; its sex can not be determined, as, un- 
fortunately, the abdomen, as I before stated, is non est, 
but from the general appearances I should suppose 
the example in question to be a 2.” Subsequently, 
Barnes and McDunnough (1918b:20) offered: “at the 
present time it is misplaced or lost . . . Fortunately, 
Strecker’s figure leaves no doubt as to the identity.” We 
have not been able to locate this specimen at the 
ANSP, and the specimen of marmorata in the box of 
supposed W. H. Edwards types at the BMNH has an 
abdomen and bears the label “Ky.,” contradicting the 
stated type locality of Yreka, California. The original 
description did not state the number of types, but it 
seems clear from the early literature accounts that only 
one specimen was known at the time. We therefore ac- 
cept the specimen figured by Strecker (1874, Plate IX, 
Fig. 6) as the holotype by monotypy for marmorata. 
The type locality of Yreka, California [USA] is erro- 
neous, since marmorata has a well-defined distribu- 
tion from the mid-Atlantic states west to Missouri and 
Arkansas (Peacock & Gall 2001). We recognize Cato- 
cala marmorata as a full species. 


Catocala minuta W. H. Edwards, 1864. Proc. Ento- 
mol. Soc. Phil. 2:512 


The original description does not state the number 
of types, but indicates “Mrs. Bridgham’s Collection. 
New York.” There are two specimens of minuta ex 
Bridgham collection at the YPM. One male bears a la- 
bel “minuta/Ed.?” in Grote’s handwriting, and another 
bears a label “undescribed A,” apparently in Edwards’ 
handwriting. These correspond to minuta and its un- 
described variety in Edwards’ description, and we 


240 


hereby designate the male with Grote’s label as LEC- 
TOTYPE for minuta (Fig. 1d). In addition to the 
above label, the lectotype bears the labels “Samuel 
Willard Bridgham/Collection/Presented Rhode Is- 
land/Audubon Society, May 1992” and “LECTO- 
TYPE/Catocala minuta/Edwards 1864/Desig. Gall & 
Hawks 2002.” The type locality is New York [USA]. 
We recognize Catocala minuta as a full species. 


Catocala nebulosa W. H. Edwards, 1864. Proc. Ento- 
mol. Soc. Phil. 2:510 


The original description does not state the number 
of types, nor the collection from which Edwards made 
his description, indicating only “Philadelphia; Wash- 
ington.” There are no nebulosa ex Bridgham collection 
at the YPM. At the ANSP collection are two old, unla- 
beled male Catocala nebulosa. However, we are not 
certain that these ANSP specimens were among those 
available to Edwards. Since the original desoaiator of 
nebulosa is diagnostic, and there has been no confu- 
sion as to the applicability of the name, we choose not 
to take any formal action. We recognize Catocala neb- 
ulosa as a full species. 


Catocala parvula W. H. Edwards, 1864. Proc. Ento- 
mol. Soc. Phil. 2:512 


The original description does not state the number 
of types, nor the collection from which Edwards made 
his description. At the ANSP collection is an old male 
bearing a label “C. minuta/v. parvula,” pinned in a dis- 
tinctive style that matches that of other Catocala la- 


beled as types at the ANSP. To clarify application of 


the name, we hereby designate this male as LECTO- 
TYPE for parvula (Fig. le). The lectotype bears the 
aforementioned label and “LECTOTYPE/Catocala 
parvula/Edwards 1864/Desig. Gall & Hawks 2002.” 
The type locality is New York [USA]. The name 
parvula is a synonym of Catocala minuta Edwards, 
and represents specimens with a large dark patch 
along the forewing inner margin. 


Catocala serena W. H. Edwards, 1864. Proc. Ento- 


mol. Soc. Phil. 2:510 


The original description does not state the number 
of types, but indicates “From Mr. Wilt’s collection, 
Philadelphia.” There are several old specimens of ser- 
ena at the ANSP. as well as at the CMNH, but we are 
not certain these were among the specimens available 
to Edwards. Since the original description of serena is 
diagnostic, and there has been no confusion as to the 
applicability of the name, we choose not to take any 

1. We recognize Catocala serena as a full 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Catocala similis W. H. Edwards, 1864. Proc. Ento- 
mol. Soc. Phil. 2:511 


The original description does not state the number 
of types, but indicates “Mrs. Bridgham’s Collection.” 
Three specimens of similis ex Bridgham collection are 
at the YPM, and to clarify application of the name we 
hereby designate a male as LECTOTYPE for similis 
(Fig. 1f). The lectotype bears the labels “Samuel 
Willard Bridgham/Collection/Presented Rhode Is- 
land/Audubon Society, May 1992” and “LECTO- 
TYPE/Catocala similis/Edwards 1864/Desig. Gall & 
Hawks 2002.” The type locality is not specified, but is 
probably Rhode Island, USA. We recognize Catocala 
similis as a full species. 


Catocala tristis W. H. Edwards, 1864. Proc. Entomol. 
Soc. Phil. 2:511 


The original description does not state the number 
of types, but indicates “From Mr. Newmans collec- 
tion, taken at Philadelphia; also Mr. Jung’s collection, 
at Hoboken, New Jersey.” There is an old male at the 
ANSP, pinned in a distinctive style matching other 
Catocala labeled as types at the ANSP. To clarify ap- 
plication of the name, we hereby designate this male 
as LECTOTYPE for tristis (Fig. 1g). The lectotype 
bears the label “LECTOTYPE/Catocala tristis/Ed- 
wards 1864/Desig. Gall & Hawks 2002.” The type lo- 
cality can not be specified further. Although Forbes 
(1954) treated tristis as a synonym of Catocala an- 
dromedae Guenée (1852), Hodges et al. (1983) re- 
elevated tristis as a northern subspecies of androme- 
dae. The differences cited for tristis are indeed not 
geographically definable, as Forbes implied, and so we 
hereby place tristis as a synonym of andromedae (RE- 
VISED SYNONYMY). 


Catocala walshti W. H. Edwards, 1864. Proc. Ento- 
mol. Soc. Phil. 2:509 


The original description does not state the number 
of types, and indicates “Taken by Mr. B. D. Walsh in 
Southern Illinois.” Grote (1873:163) stated that the 
species “is still unknown to me. I believe the types per- 
ished in the Chicago fire,” and Smith (1893) appar- 
ently followed him in stating the types were destroyed. 
However, the 14 February 1879 minutes of the Pro- 
ceedings of the Monthly Meetings of the Entomological 
Section of the Academy of Natural Sciences, Philadel- 
phia indicate (Anonymous 1879:iii): “Mr. Cresson ex- 
hibited the type specimens of Catocala editha Ed- 
wards, and Catocala walshii Edwards, which had been 
sent to him by Mr. William H. Edwards, for the collec- 
tion of the American Entomological Society.” There 


VOLUME 56, NUMBER 4 


are several old specimens at the ANSP and CMNH la- 
beled as walshii, but we can not say for certain which 
of these might be the Cresson specimen(s). 

The type locality of southern Illinois suggests wal- 
shii refers to Catocala junctura Walker, the only large 
Catocala species in that geographic region that 
matches the original description, and walshii has been 
treated as a synonym of juwnctura since Barnes and Mc- 
Dunnough (1918b:29). However, because (a) we have 
been unable to locate a type, (b) other published type 
localities of Edwards are erroneous, and (c) a wil- 
low/poplar feeding taxon is involved, we feel the name 
walshii requires explicit clarification. We therefore 
designate a male from among the aforementioned 
specimens at the ANSP as NEOTYPE for walshii (Fig. 
lh). The neotype bears the label “Catocala/walshii” 
and “NEOTYPE/Catocala walshii/Edwards 1864/De- 
sig. Gall & Hawks 2002.” The type locality is southern 
Illinois [USA]. The name walshii is a synonym of Cato- 
cala junctura Walker. 


AUGUSTUS R. GROTE 


Augustus Radcliffe Grote was one of the foremost 
authorities on Nearctic moths in the 19th century. He 
published profusely throughout the Noctuidae, and 
authored 45 Nearctic Catocala names, more than any 
other worker. His earliest Catocala descriptions were 
published in 1864 and 1866 (with Coleman T. Robin- 
son), and in 1872, Grote presented the first revision of 
the entire Nearctic Catocala fauna in the Transactions 
of the American Entomological Society. This revision 
would doubtless have been even more comprehensive, 
if it were not for the fact that Grote apparently lost the 
original manuscript for it. 

Grote described new species from specimens in a 
wide array of personal and institutional collections, 
and hence his types are known to be widely scattered. 
The BMNH purchased most of Grote’s collection in 
1881/1882, and this is the principal repository of his 
type material. For Grote Catocala types, the ANSP is 
as important as the BMNH, as Grote remarked in his 
1872 revision that “most of the material I have used is 
in the collection of the American Entomological So- 
ciety, together with Mr. Edwards’ specimens kindly 
loaned me for examination.” 

Although we located types for most Grote names 
fairly readily, we encountered some difficulties track- 
ing down types from his earliest publications, notably 
the Catocala he authored with C. T. Robinson. This is 
hardly a novel situation, as lepidopteran systematists 
have speculated for some time on the whereabouts of 
apparently lost Grote and Robinson types. Beuten- 
miiller (1892) provides an accounting of Lepidoptera 


types known to be at the AMNH from the Grote and 
Robinson collection, but no Catocala are included in 
that list. Smith’s (1893:12) discussion is instructive: 
“Mr. Grote writes concerning the work by Mr. Robin- 
son and himself: ‘The collection on which these and all 
our other joint entomological writings were based, is 
now in the possession of the American Entomological 
Society. If that was true in 1872, it certainly is not so 
now, because but a very few of the species are at pres- 
ent in that collection. Of these, a very small proportion 
only have a written label. Some have a little printed 
‘type’ label; but nothing to indicate of what it is the 
type. In at least two cases I found these ‘type’ speci- 
mens under names with which they had no possible 
relationship. Fortunately the excellent figures given in 
illustration of the papers referred to, make it possible 
to ascertain the names of which the specimens are typ- 
ical. But many species have disappeared, leaving no 
trace; in whose hands they are at present I have been 
unable to ascertain.” 

Grote moved to Germany in 1884 after the sale of 
his collection to the BMNH, living first in Bremen and 
then later in Hildesheim until his death in 1903. He 
was actively collecting while at Hildesheim (e.g., Grote 
1897, Stein 1999), and Hom and Kahle (1935:97) indi- 
cated that Grote’s material was also deposited “ex 
parte 1893 an Mus. Bremen.” Since it seemed possible 
that the missing Grote and Robinson material to which 
Smith (1893) referred could be in Germany, we visited 
the Museum fur Naturkunde of Humboldt University 
in Berlin (ZMHU), the Ubersee-Museum in Bremen 
and the Roemer- und Pelizaeus Museum in Hildes- 
heim in April 2000 to search for types. We were able to 
review the entire lepidopteran holdings of the Roemer 
Museum, all the Nearctic Lepidoptera at the Ubersee 
Museum, but only the catocalines at ZMHU. The 
ZMHU has 24 drawers of Catocala, and it contains the 
types for two Grote Catocala species that he explicitly 
indicated were in that institution, as well as a number 
of specimens that he and Robinson determined. The 
Ubersee Museum collection has 4 drawers of Grote’s 
Nearctic Lepidoptera, two drawers of which are exclu- 
sively Catocala, but no types. The Roemer Museum 
collection, presently housed at the Biology Depart- 
ment of Hildesheim University, has 13 drawers of 
Grote’s Nearctic Lepidoptera, several of which contain 
Catocala, including the type of Catocala moderna 
Grote (as stated in the original description). The Roe- 
mer also has the Palearctic Lepidoptera that Grote as- 
sembled from Hildesheim and surrounding areas. 

We have also examined the Lepidoptera collection 
at the Buffalo Museum of Science, which contains the 
holdings from the Buffalo Society of Natural History, 


with which Grote was affiliated during the 1870's when 
he lived in Buffalo. We found no Grote Catocala types 
there. Arnett et al. (1993:209) state that “major por- 
tions of [Grote’s] collection were given to the Museum 
of Natural History (BMNH) and to the Museum of 
Natural History of Sao Paulo,” and thus Brazil is the 
only other potential repository for Grote and Robinson 
material of which we are aware. Insofar as Catocala 
are concerned, the Museu de Zoologia in Sao Paulo 
currently has only a small number of specimens, rep- 
resenting common Nearctic and Palearctic species. 
Among the Sao Paulo specimens are one cara Guenée, 
one concumbens Walker, and one ilia (Cramer), all 
bearing Grote determination labels in his handwriting 
(C. R. F. Brandao pers. com. 1999). 

At this juncture we believe that the missing lepi- 
dopteran types of Grote and Robinson are probably 
not extant in Germany, and we are more confident still 
that this is the case for their Catocala types. In sum, 
then, definitive and probable type material for the 
Catocala authored by Grote, as well as Grote and 
Robinson, exist in the collections of the ANSP, 
BMNH, AMNH, ZMHU, CMNH and at Hildesheim 
University. The presence at CMNH of Grote Catocala 
is the result of an extensive exchange of material be- 
tween ANSP and CMNH in the 1960's, that sent Lep- 
idoptera from ANSP to CMNH, and Orthoptera from 
CMNH to ANSP (J. Rawlins pers. com. 1999). At 
CMNH, these former ANSP specimens are identifi- 
able by labels indicating they were part of CMNH ac- 
cession 20359. 

In most instances, Grote types in the aforemen- 
tioned institutional collections bear one or more labels 
indicating they are types, although this is not univer- 
sally so (e.g., the BMNH holotype of Catocala in- 
nubens var. flavidalis Grote, mentioned by Beuten- 
miiller 1903b). The label situation at the BMNH, 
where the bulk of Grote’s collection resides, merits 
further elaboration. Many of Grote’s Catocala types at 
the BMNH have apparently been identified as such by 
A. G. Butler (or his curatorial contemporaries), whose 
habit was to write the taxon name and the word “type” 
on the reverse of the attached BMNH accession label 
(M. Honey pers. com. 1991). Typically, these types also 
bear a red-bordered label with the taxon name fol- 
lowed by the word “type,” handwritten by Grote. Most 
of Grote’s other Catocala specimens at the BMNH 
bear no type labels, but a small number of these other 
specimens bear blue-bordered labels with only the 
taxon name, again in Grote’s handwriting (in at least 
some other noctuid groups, Grote’s habit appears to 

| a red-bordered label when one type 


i} ) ii tO us 
ed, and a blue-bordered label when more than 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


one existed; E. Quinter pers. com. 2001). The blue- 
bordered labels often appear on single exemplars of 
Catocala named by Grote in the 1870's, and by Grote 
and Robinson in 1866 (one such specimen of scintil- 
lans Grote & Robinson is also labeled as type by But- 
ler, the only blue-bordered Grote & Robinson taxon so 
labeled at the BMNH). A few specimens with the 
blue-bordered Tabels also bear the red-bordered type 
labels: for example, the type of sinwosa Grote, named 
in 1879, and the types of coelebs Grote and semirelicta 
Grote, both of which were named in 1874. Based on 
the distribution of these red and blue-bordered labels 
among the BMNH Catocala, we believe that speci- 
mens bearing such labels were among those on which 
Grote based his type concepts, and so we treat them 
all as available type material. 

We also note here that the two published listings of 
Nearctic Catocala types at the BMNH (Beutenmiiller 
1903a, Hampson 1913; both were prepared by Hamp- 
son) are not concordant. The former tabulates 27 taxa, 
the latter 31 taxa. There are 25 taxa common to both 
the 1903 and 1913 listings. Six taxa appear only in the 
1913 listing (beaniana Grote, crataegi Saunders, elec- 
tilis Walker, nurus Walker, selecta Walker, subnata 
Grote), and two appear only in the 1903 listing 
(bunkeri Grote, sordida Grote). In addition, the types 
of albomacula Butler, flavidalis and scintillans Grote 
& Robinson are omitted from both the 1903 and 1913 
listings, despite the fact that the main collection at the 
BMNH has specimens of each that bear type labels 
written by Butler. Because of these omissions, the 
dropping of bunkeri and sordida between 1903 and 
1913, and the inconsistent tabulation of the Walker 
types (these had long since been established e.g., see 
Grote & Robinson 1868), we view with some reserva- 
tion Hampson’s compilations of type specimens of 
Nearctic Catocala at the BMNH. 

Thus, our approach for Grote Catocala is to give 
precedence first to specimens having the most appro- 
priate label data in the most appropriate collections, 
and then to the BMNH and ANSP/CMNH collec- 
tions, in that order. However, for taxa described in 
Grote’s (1872) revision, the ANSP/CMNH is given ini- 
tial precedence. For taxa for which we failed to locate 
a labeled type, we choose BMNH specimens with 
blue-bordered Grote labels, if possible. Although the 
published type localities for Grote names usually 
match data on the specimen labels, this is not univer- 
sally so. The most common mismatches involve the 
handwritten locality data that were added subse- 
quently to the printed “Grote Coll./81-116” accession 
labels by BMNH curatorial staff. In such instances, we 
give precedence to Grote’s published type localities. 


VOLUME 56, NUMBER 4 


243 


Fic. 2. Type specimens of moths in the genus Catocala Schrank. A. R. Grote types (authorship is Grote unless otherwise noted). a, lecto- 
type, abbreviatella. b, lectotype, alabamae. ec, lectotype, anna. d, lectotype, badia Grote & Robinson. e, lectotype, coccinata. f, lectotype, che- 
lidonia. g, lectotype, dulciola. h, lectotype, formula Grote & Robinson. i, lectotype, communis. j, lectotype, flebilis. 0.9-1.0 x natural size. 

& /p YE YE j YPe:, 


Available Names 


Catocala abbreviatella Grote, 1872. Trans. Amer. 
Entomol. Soc. 4:14 


The original description states “d° . . . Two speci- 
mens from Texas (Belfrage).” There are only two ab- 
breviatella at the ANSP, one male and one female, 


both old and bearing no labels but pinned in a similar 
manner. In Drawer 25 at the BMNH is one male bear- 
ing a conflicting locality label of “Kansas.” The pair at 
the ANSP appear to be the syntypes, and to clarify ap- 
plication of the name we hereby designate the male as 
LECTOTYPE for abbreviatella (Fig. 2a). The lecto- 
type bears the label “LECTOTYPE/Catocala abbrevi- 


atella/Grote 1872/Desig. Gall & Hawks 2002.” The 
type locality is Texas [USA]. We recognize Catocala 
abbreviatella as a full species. 


Catocala alabamae Grote, 1875b. Proc. Acad. Nat. 

Sci. Phil. 27:427 

The original description does not state the number 
of types. In Drawer 29a at the BMNH is a female la- 
beled as type by Grote, bearing a red BMNH type 
disc. To clarify application of the name we hereby des- 
ignate it as LECTOTYPE for alabamae (Fig. 2b). The 
lectotype bears the labels “Ala.”, “Alabama/Grote 
Coll./81-116.”, “Alabama/81-116 [and on the reverse: | 
Catocala/alabamae/Type Grote”, “Catocala alaba- 
mae/Type Grote”, “LECTOTYPE/Catocala alaba- 
mae/Grote 1875/Desig. Gall & Hawks 2002.” The type 
locality is Demopolis, [Marengo County,] Ala[bama, 
USA]. We recognize Catocala alabamae as a full 
species. 


Catocala adoptiva Grote, 1874c. Trans. Amer. 
Entomol. Soc. 5:96. 


The male lectotype is at the MCZ, and was desig- 
nated by Gall (1990:121). The type locality is Dallas 
Co[unty], Texas, [USA]. We recognize Catocala adop- 
tiva as a synonym of Catocala delilah Strecker. 


Catocala angusi Grote, 1876a. Can. Entomol. 8:229 


We consider Beutenmiiller’s (1907:150) action to 
constitute a valid lectotype designation for angusi un- 
der Article 74 of the Code. The male lectotype is at the 
AMNH. The type locality is West Farms, New York 
City [Bronx County, USA], fide Beutenmiiller’s state- 
ments. We recognize Catocala angusi as a full species. 


Catocala anna Grote, 1874c. Trans. Amer. Entomol. 
Soc. 5:96 


The original description states “Texas. (Boll. No. 
104).” In Drawer 25 at the BMNH is a male labeled as 
anna by Grote, but not as type. An identically labeled 
male is at the ANSP. A total of five other new Catocala 
were described by Grote in the article in which anna 
appeared, and the types for these five other taxa are at 

the BMNH. We thus give precedence to the BMNH 
specimen in Drawer 25, and to clarify application of 
the name designate it as LECTOTYPE for anna (Fig. 
2c). The lectotype bears the labels “anna Grote”, “U. S. 
America/Grote Coll./81-116.”, “United States/81-116 
[and on the reverse:] Catocala/anna/Grote”, “Cato- 
cala/anna/Grote’, “1 ECTOTYPE/Catoc ala anna/Grote 
1874/Desig. Gall & Hawks 2002.” The type locality is 
Texas [USA]. The name anna is a synonym of Catocala 
amestris Strecker. 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Catocala arizonae Grote, 1873. Can. Entomol. 5:163 


The original description states “I have received this 
species from Professor Townend Glover, of the Agri- 
cultural Department at Washington. It is labelled “Bor- 
ders of Arizona and New Mexico.—Dr. Palmer.” This 
male, the holotype by monotypy, is at the ANSP 
(ANSP Type #7659). The type locality is the border of 
Arizona and New Mexico [USA]. The type of arizonae 
is a specimen of Catocala junctura Walker, and we 
hereby place arizonae as a synonym of junctura (NEW 
SYNONYMY). 


Catocala badia Grote & Robinson, 1866. Proc. 
Entomol. Soc. Phil. 6:22 


The original description states “Numerous coinci- 
dent specimens from Massachusetts, Rhode Island, 
New York and Pennsylvania examined.” Among five 
old specimens at the ANSP is a female bearing the ma- 
chine-printed labels “Mass.” and “Coll. of Acd./Nat. 
Sci. Phila.” A male in Drawer 3a at the BMNH bears a 
blue-bordered Grote label. We give precedence to the 
BMNH specimen, and to leas application of the 
name we hereby designate this male as LECTOTYPE 
for badia (Fig. 2d). The lectotype bears the labels “U 
S. America/Grote Coll./81-116,” “United States 81-116 
[and on the reverse] Catocala/badia/Gr. + Rob.,” 
‘Catocala’ ‘Schrank/badiay Gs Ree Le GnOmMrE 
Catocala badia/Grote & Robinson 1866/Desig. Gall & 
Hawks 2002.” The type locality cannot be restricted 
further. We recognize Catocala badia as a full species. 


Catocala beaniana Grote, 1878. Can. Entomol. 
10:195 


The original description states “Sent me by Mr. 
Thomas A. Bean, under the number 574, from Illinois.” 
This female, the holotype by monotypy, is in Drawer 18 
at the BMNH. The type locality is Illinois [USA]. The 
name beaniana is a synonym of Catocala meskei Grote. 


Catocala chelidonia Grote, 1881c. Papilio 1:159 


The original description states “2 . . . Numerous 
specimens of this very distinct form in the collection 
[Neumoegen] before me.” A male type is at the 
USNM, and to clarify application of the name we 
hereby designate it as LECTOTYPE for chelidonia 
(Fig. 2f). The lectotype bears the labels “Prescott/ 
Arizona.’, “Type No./33987/U.S.N.M.”, “TYPICUM/ 
SPECIMEN”, “Catocala Type/chelidonia/Grote’, 
“LECTOTYPE/Catocala chelidonia/Grote 1881/De- 
sig. Gall & Hawks 2002.” The type locality is Prescott, 
[Yavapai County,] Arizona [USA]. We recognize Cato- 
cala chelidonia as a full species. 


VOLUME 56, NUMBER 4 


Catocala clintoni Grote, 1864a. Proc. Entomol. Soc. 
Phil. 3:89 


The original description states “Hab. Eastern States. 
A ° specimen in good condition given me by Mr. Wm. 
A. Nason, and now in the Coll. Ent. Soc. Philadel- 
phia.” There are four clintoni in the ANSP collection, 
two with locality labels indicating Kansas and two indi- 
cating Texas, none of which match the locality of 
“Eastern States.” In Drawer 23 at the BMNH is a fe- 
male clintoni from New York, ex Grote collection. An 
old female clintoni at the CMNH bears a label indicat- 
ing it came from accession 20359. We know that at 
least one type of praeclara Grote & Robinson ended 
up at CMNH as part of the CMNH/ANSP exchange 
on accession 20359. Because Hampson (1913) and 
Beutenmiiller (1903a) omit clintoni from their lists of 
Nearctic Catocala types at the BMNH, and because 
the original description calls for the ANSP, we believe 
that the CMNH specimen is more probably the holo- 
type by monotypy. The type locality is Eastern States 
[USA]. We recognize Catocala clintoni as a full 
species. 


Catocala coccinata Grote, 1872. Trans. Amer. 
Entomol. Soc. 4:6 


The original description states “d° ... My specimens 
are from Pennsylvania.” A male type is at the ANSP, 
and to clarify application of the name we hereby des- 
ignate it as LECTOTYPE for coccinata (Fig. 2e). The 
lectotype bears the labels “Penn.”, “TYPE No. 
7663/Catocala/coccinata/A. R. Grote”, “C. cocci- 
nata/Grote/A.R.G.”, “LECTOTYPE/Catocala cocci- 
nata/Grote 1872/Desig. Gall & Hawks 2002.” The type 
locality is Pennsylvania [USA]. We recognize Catocala 
coccinata as a full species. 


Catocala coelebs Grote, 1874c. Trans. Amer. 
Entomol. Soc. 5:96 


The original description states “St. Catherines, Au- 
gust 18, No. 182, Geo. Norman Esq.” This female, the 
holotype by monotypy, is in Drawer 4 at the BMNH. 
The type locality is S[ain]t Catherines, [Regional Mu- 
nicipality of Niagara, Ontario, Canada]. 

Catocala coelebs and Catocala badia Grote & 
Robinson have been treated for some time as closely 
related, distinct species. Largely monomorphic 
populations referable to badia occur along the east- 
ern seaboard from south central Massachusettes 
through New Jersey and Pennsylvania; similarly 
monomorphic populations referable to coelebs occur 
from the Maritime Provinces of Canada through 
Maine and northern New England, and westward to 


northern Michigan, Wisconsin and the Prairie 
Provinces of Canada. However, the geographic 
ranges of these “pure” badia and “pure” coelebs 
populations are essentially parapatric—a small zone 
of overlap exists in New Hampshire, southern 
Maine, and northern coastal Massachusetts in which 
specimens with intermediate forewing patterns oc- 
cur. These have been named badia form phoebe 
Hulst. 

We have examined large series of badia and coelebs 
from the entire geographic range of both taxa, particu- 
larly in the zone of overlap in New England, and find 
no consistent way to distinguish material in the overlap 
area. Although we have reared a number of broods of 
“pure” badia from Connecticut and southern Massa- 
chusetts, and “pure” coelebs from northern Maine and 
Michigan, eggs from a female coelebs taken in south- 
ern York County, Maine produced progeny referable 
to both coelebs (several dozen adults) and phoebe 
(three adults) and intergrades. We have also found no 
constant differences between badia and coelebs in 
their larvae or the scanning electron micrograph pat- 
terns of their eggs. However, the larvae and eggs of ba- 
dia/coelebs are separable from those of the two other 
closely related Nearctic Myricaceae-feeding species, 
antinympha (Hiibner) and muliercula Guenée; in 
turn, the larvae of antinympha and muliercula are also 
separable from one another. 

Accordingly, we feel the observed geographic varia- 
tion in badia and coelebs is more indicative of two dis- 
tinct subspecies, and we hereby place the name 
coelebs as a subspecies of Catocala badia, new status. 
Grote, who described both badia and coelebs, had ar- 
rived at an essentially similar conclusion (1883a:23): 
“But a sight of many specimens leads me to believe 
that the true C. coelebs, with black fore wings, grades 
into badia, and is only to be regarded as a variety of it. 
But this could not have been predicated of the type [of 
coelebs|. The two extremes are exceedingly distinct 
and different looking.” 


Catocala communis Grote, 1872. Trans. Amer. 
Entomol. Soc. 4:9 


In a discussion of Catocala neogama J. E. Smith, 
Grote stated (1872:9) “I do not believe this [neogama 
sensu Guenée] to be the Phalaena neogama of Smith. 
In the Berlin Museum I have named this species Cato- 
cala communis, while two specimens from Texas 
therein contained differed from the present by their 
brighter colored secondaries, and otherwise more 
nearly resembled Abbot's figure [of neogama J. E. 
Smith].” A male type is in Drawer 155 at the Museum 
fur Naturkunde of Humboldt University in Berlin 


(ZMHU), and to clarify application of the name we 
hereby designate it as LECTOTYPE for communis 
(Fig. 2i). The lectotype bears the labels “10918”, “com- 
munis/Grote & Rob./neogama Guen/(non Sm.- 
Abb.)/Carolina [unintelligible], “7158”, “LECTO- 
TYPE/Catocala communis/Grote 1872/Desig. Gall & 
Hawks 2002.” The type locality is hereby restricted to 
|North or South] Carolina, [USA] on the basis of the 
lectotype label. The name communis is a synonym of 
Catocala neogama J. E. Smith. 


Catocala dulciola Grote, 1881a. Papilio 1:5 


The original description states “Five or six examples, 
all alike, have been collected by Mr. Pilate . . . The fe- 
male type before me expands 44 millimetres.” A fe- 
male type is at the AMNH, and to clarify application of 
the name we hereby designate it as LECTOTYPE for 
dulciola (Fig. 2g). The lectotype bears the labels 
‘Catocala 2° Type/dulciola Grote”, “No. 11742 H. 
Edw.”, “Type No. AMNH”, “Catocala/dulciola/Grote’, 
“LECTOTYPE/Catocala dulciola/Grote 1881/Desig. 

Gall & Hawks 2002.” The type locality is the vicinty of 

Dayton, [Montgomery County], Ohio [USA]. We rec- 
ognize Catocala dulciola as a full species. 


Catocala flebilis Grote, 1872. Trans. Amer. Entomol. 
Soc. 4:4 


The original description states “Several specimens 
examined from Pennsylvania.” A male type is at the 
ANSP, and to clarify application of the name we 
hereby designate it as LECTOTYPE for flebilis (Fig. 

2)). The lectotype bears the labels “Penn.”, “TYPE No. 
ata Canta fred IR, Grow”, “C, Aelita Grate! 
A.R.G./Type”, “LECTOTYPE/Catocala flebilis/Grote 
1872/Desig. Gall & Hawks 2002.” The type locality is 
Pennsylvania [USA]. We recognize Catocala flebilis as 
a full species. 


Catocala formula Grote & Robinson, 1866. Proc. En- 
tomol. Soc. Phil. 6:27 


The original description states “A number of speci- 
mens of both sexes examined. Habitat—New York State, 
Rhode Island. Coll. Ent. Soc. Phila.; our own Collection 
and that of Mrs. S. W. Bridgham, N. Y.” A male type is at 
the ANSP, and to clarify application of the name we 
hereby designate it as LECTOTYPE for formula (Fig, 
2h). The lectotype bears the labels “N. Y.”, “TYPE No. 
7662/Catocala/formula/A. R. Grote & Rob.”, “GC. formula 
G + R’, “LECTOTYPE/Catocala formula/Grote & 
Robinson 1866/Desig. Gall & Hawks 2002.” The type lo- 


cé ality is he sreby restricted to New York [USA] on the ba- 
ctotype locality label. The name formula is a 
synonym of Catocala similis Edwards. 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Catocala fratercula Grote & Robinson, 1866. Proc. 
Entomol. Soc. Phil. 6:24 


The original description lists “Habitat——New York 
State. Rhode Island. Several (¢ and 2) specimens ex- 
amined.” A male type is at the ANSP, and to clarify ap- 
plication of the name we hereby designate it as 
LECTOTYPE for fratercula (Fig. 3a). The lectotype 
bears the labels “N. Y.”, “TYPE No. 7661/Cato- 
cala/fratercula/A. R. Grote & Rob.”, “C. fratercula/G + 
R/TYPE”, “LECTOTYPE/Catocala fratercula/Grote 
& Robinson 1866/Desig. Gall & Hawks 2002.” The 
type locality is hereby restricted to New York [USA] on 
the basis of the lectotype locality label. Although 
Forbes (1954) placed fratercula as a synonym of Cato- 
cala micronympha Guenée, Hodges et al. (1983) re- 
elevated fratercula as a northern subspecies. However, 


fratercula is indeed merely one of many infrapopula- 


tional variants found over a wide geographic range in 
this species, and so we return fratercula as a synonym 
of micronympha (REVISED SYNONYMY). 


Catocala frederici Grote, 1872. Trans. Amer. Ento- 


mol. Soc. 4:14 


The original description lists “Iwo specimens from 
Texas (Friedrich) examined in the Berlin Museum.” 
These two female syntypes are in Drawer 157 at the 
Museum fur Naturkunde of Humboldt University in 
Berlin (ZMHU), and to clarify application of the name 
we hereby designate the first female as LECTOTYPE 
for frederici (Fig. 3b). The lectotype bears the labels 
“10923”, “frederici/Grote & Rob./Texas. Friedrich”, 
“LECTOTYPE/Catocala frederici/Grote 1872/Desig. 
Gall & Hawks 2002.” The type locality is Texas [USA]. 
We recognize Catocala frederici as a full species. 


Catocala guenei Grote, 1887. Can. Entomol. 19:115. 


The name guenei is the second published and hence 
unnecessary replacement name for Catocala viduata 
Guenée (1852). The first published replacement name 
for viduata is Catocala maestosa Hulst (1884). Since 
guenei is a replacement name, its type locality is the 
same as for Guenée’s viduata, namely Georgia, [USA]. 


Catocala habilis Grote, 1872. Trans. Amer. Entomol. 
Soc. 4:11 
The original description states “A number of coinci- 
dent specimens examined from Pennsylvania.” A male 
type is at the ANSP, and to clarify application of the 
name we hereby designate it as LECTOTYPE for ha- 
bilis (Fig. 3d). The lectotype bears the labels “Penn.”, 
“TYPE No. 7664/Catocala/habilis/A. R. Grote”, “Cato- 
cala/habilis/Grote/TYPE”, “LECTOTYPE/Catocala ha- 
bilis/Grote 1872/Desig. Gall & Hawks 2002.” The type 


VOLUME 56, NUMBER 4 


bo 
oS 
~l 


Fic. 3. Type specimens of moths in the genus Catocala Schrank. A. R. Grote types (authorship is Grote unless otherwise noted). a, lecto- 
type, fratercula Grote & Robinson. b, lectotype, frederici. e, lectotype, levettei. d, lectotype, habilis. e, lectotype, meskei. f, lectotype, lincella. 
g, lectotype, mira. h, lectotype, praeclara Grote & Robinson. i, lectotype, phalanga. j, lectotype, piatrix. 0.9-1.0 x natural size. 


locality is Pennsylvania [USA]. We recognize Catocala 
habilis as a full species. 


Catocala levettei Grote, 1874c. Trans. Amer. Entomol. 
Soe. 5:95 


The original description does not state the number 
of types. A male labelled as type by Grote is in Drawer 


1 at the BMNH, bearing a red BMNH type disc. The 
specimen label indicates Ohio, but the original de- 
scription calls for Indianapolis. Hampson (1913) and 
Beutenmiiller (1903a) stated the type of levettei was at 
the BMNH, and we have no other reason other than 
this probable labeling error to believe this BMNH 
male is not an available syntype. To clarify application 


of the name we hereby designate it as LECTOTYPE 
for levettei (Fig. 3c). The lectotype bears the labels 
“Ohio./Grote Coll./81-116.”, “Ohio/81-116 [and on the 
reverse: | Catocala/levettei/Type Grote”, “Catocala/lev- 
ettei/Type. Grote”, “LECTOTYPE/Catocala levet- 
tei/Grote 1874/Desig. Gall & Hawks 2002.” We give 
precedence to Grote’s original description and retain 
the type locality as Indianapolis, [Marion County, In- 
diana, USA]. The name levettei is a synonym of Cato- 
cala judith Strecker. 


Catocala lineella Grote, 1872. Trans. Amer. Entomol. 
Soc. 4:18 


The systematics of the “amica complex’ of small 


Nearctic Catocala species has been the recent focus of 


breeding and field work by D. F. Schweitzer and our- 
selves. We know of four species, three of which have 
available names, and one undescribed sibling species 
that lacks an available name. Catocala jair Strecker is 
the rarest, most distinctive, and most geographically 
restricted of the four species. The other three are 
widespread, with Catocala amica (Hiibner) and Cato- 
cala lineella Grote being the most frequently encoun- 
tered in most areas of eastern North America. Cato- 
cala lineella had been treated as a subspecies or 
synonym of Catocala amica during the 20th century, 
but it was restored to the rank of full species by Gall 
(1990). 

The usage and applicability of amica and jair have 
been fixed in the literature for some time, and al- 
though the usage of lineella has been modestly stable, 
Barnes and Mc Dwmnengh (1918b) were unaware of a 
type. The original description of lineella indicates “a 
number of coincident specimens.” In Drawer 34 at the 
BMNH is a male with a blue-bordered Grote label. To 
clarify application of the name we hereby designate it 
as LECTOTYPE for lineella (Fig. 3f). The lectotype 
bears the labels “E. U. S. America/Grote Coll./81- 
116.”, “19/6”, “8.”, “E. States/81-116 [and on the re- 
verse:| Allotria/lineella/Grote”, “Catocala /Schk./li- 
neella/Grote”, “~LECTOTYPE/Catocala lineella/Grote 
1872/Desig. Gall & Hawks 2002.” Grote listed the type 
locality for lineella as “Same localities as C. an- 
drophila,” which was “Eastern States, southward.” We 
recognize Catocala lineella as a full species. 


Catocala meskei Grote, 1873. Can. Entomol. 5:161 


The original description states “Lent me by Mr. O. 
Meske, after whom I name the species, from near 


Albany, N. Y., and who writes me that it has been taken 
in considerable numbers by a collector in that vicinity.” 
Although ‘sa single specimen, the number 


of types is not explicitly stated. To clarify application of 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


the name we hereby designate the female at the 
USNM as LECTOTYPE for meskei (Fig. 3e). The 
lectotype bears the labels “August/1873.”, “Cato- 
cala/meskei/Grote/Type”, “Type/No. 305/U.S.N.M.” 
“Collection/O. Meske’, “LECTOTYPE/Catocala 
meskei/Grote 1873/Desig. Gall & Hawks 2002.” The 
type locality is Albany, [Albany County,] N[ew] Y[ork, 
USA]. We recognize Catocala meskei as a full species. 


Catocala mira Grote, 1876a. Can. Entomol. 8:230 


The original description does not state the number 
of types. In Drawer 29 at the BMNH is a female with 
a red-bordered Grote type label, bearing a red BMNH 
type disc. To clarify application of the name we hereby 
designate it as LECTOTYPE for mira (Fig. 3g). The 
lectotype bears the labels “388”, “U. S. America/Grote 
Coll./81-116.”, “United States/81-116 [and on the re- 
verse:| Catocala/mira/Type Grote”, “C. mira/Grote/ 
Type’, “LECTOTYPE/Catocala mira/Grote 1876/De- 
sig. Gall & Hawks 2002.” The type locality is hereby re- 
stricted to the USA on the basis of the lectotype local- 
ity label. We recognize Catocala mira as a full species. 


Catocala moderna Grote, 1900. Can. Entomol. 
32:191 


The original description states “Accompanying the 
fresh type [of moderna] are examples of C. viduata, 
Guen.” The male holotype by monotypy is in Drawer 
33 of Cabinet “Neu 3” at the Roemer- und Pelizaeus 
Museum (RUPM) in Hildesheim, Germany. The type 
locality is Dolores, Texas [USA]. The name moderna is 
a synonym of Catocala maestosa Hulst. 


Catocala phalanga Grote, 1864a. Proc. Entomol. Soc. 
Phil. 3:86 


The original description states “Coll. Ent. Soc. 
Philadelphia . . . Possibly a variety of Catocala 
palaeogama, but, as several coincident specimens have 
occurred, it may be distinct.” There are several old, 
unlabeled specimens of phalanga at the ANSP, includ- 
ing one pinned in a style like that of other Grote types 
at ANSP. In Drawer 4 at the BMNH is a male bearing 
a blue-bordered Grote label. We give precedence to 
the BMNH male labeled by Grote, and to clarify ap- 
plication of the name hereby designate it as LECTO- 
TYPE for phalanga (Fig. 3i). The lectotype bears the 
labels “U. S. America/Grote Coll./81-116.”, “var. pha- 
langa/Grote”, “United States/81-116 [and on the re- 
verse:| “C. palaeogama/var. phalanga/Grote”, “LEC- 
TOTYPE/Catocala phalanga/Grote 1864/Desig. Gall 
& Hawks 2002.” The type locality is Middle States 
[USA]. The name phalanga is a synonym of Catocala 
palaeogama Guenée, and represents specimens with a 


VOLUME 56, NUMBER 4 


prominent black postmedian line and black basal 
patch on the forewing. 


Catocala piatrix Grote, 1864a. Proc. Entomol. Soc. 

Phil. 3:88 

The original description states “Coll. Ent. Soc. 
Philadelphia,” but does not indicate the number of 
specimens. In Drawer 5 at the BMNH is a male bearing 
a blue-bordered Grote label. There are several old 
males at the ANSP, but none labeled by Grote. A male 
now at the CMNH, ex ANSP via accession 20359, is la- 
beled as piatrix by Grote, and bears a machined locality 
label like that on other ANSP Catocala types. We give 
precedence to the CMNH male, and to clarify applica- 
tion of the name hereby designate it as LECTOTYPE 
for piatrix (Fig. 3j). The lectotype bears the labels 
SING “Catocala/piatrix/Gr/A.R.G.”, “Exch. A.N.S.P/C. 
M. Acc. 20359”, “LECTOTYPE/Catocala piatrix/Grote 
1864/Desig. Gall & Hawks 2002.” The type locality is 
hereby restricted to N[ew] Y[ork, USA] on the basis of 
the lectotype locality label. We recognize Catocala pia- 
trix as a full species. 


Catocala ponderosa Grote & Robinson, 1866. Proc. 
Entomol. Soc. Phil. 6:23 


The original description states “Habitat.—Illinois, 
New York, Pennsylvania. Coll. Ent. Soc. Philadelphia. 
Several specimens examined.” Two unlabeled male neb- 
ulosa are at the ANSP. In Drawer 4a at the BMNH is a 
female bearing a blue-bordered Grote label. We give 
precedence to the BMNH female labeled by Grote, and 
to clarify application of the name hereby designate it as 
LECTOTYPE for ponderosa (Fig. 4a). The lectotype 
bears the labels “6”, “U.S. America/Grote Coll./81-116”, 
“Catocala/ponderosa/G + R’, “United States/81-116 
[and on the reverse:] Catocala/ponderosa/Gr. + Rob.”, 
“LECTOTYPE/Catocala ponderosa/Grote & Robinson 
1866/Desig. Gall & Hawks 2002.” The type locality can 
not be restricted further. The name ponderosa is a syn- 
onym of Catocala nebulosa. 


Catocala praeclara Grote & Robinson, 1866. Proc. 
Entomol. Soc. Phil. 6:25 


The original description states “Our Collection and 
that of Ent. Soc. Phil., numerous (d and 2) coincident 
specimens examined.” At the CMNH is a female 
praeclara, ex ANSP via accession 20359, bearing a hand- 
written label in pencil stating “presumed TYPE.” This 
penciled label is in the same handwriting as penciled 
“TYPE” designations on other types at the ANSP. To 
clarify application of the name we hereby designate this 
CMNH female as LECTOTYPE for praeclara (Fig. 3h). 
In addition to the above label, the lectotype bears the la- 


bels “N. Y.”, “praeclara”, “Exch. A.N.S.P/C. M. Acc. 
20359", “LECTOTYPE/Catocala praeclara/Grote & 
Robinson 1866/Desig. Gall & Hawks 2002.” The type lo- 
cality is New York [USA]. We recognize Catocala 
praeclara as a full species. 


Catocala residua Grote, 1874a. Proc. Boston Soc. 
Nat. Hist. 16:242 


The original description lists “Hab. New York (in 
Coll. Buf. Soc. Nat. Sciences); Canada (J. Petit). Three 
coincident specimens examined,” and Smith (1893:354) 
states “The type of residua is in the American Museum 
of Natural History from the Angus Collection.” This 
female is at the AMNH, and to clarify application of the 
name we hereby designate it as LECTOTYPE for 
residua (Fig. 4b). The lectotype bears the labels “No. 
486/Coll. J. Angus/West Farms,/New York City.”, 
“Type/No./A.M.N.H.”, “C. residua/Grote/var.”, “LEC- 
TOTYPE/Catocala residua/Grote 1874/Desig. Gall & 
Hawks 2002.” Because Angus often affixed a printed 
West Farms label as a “return address,” rather than as a 
locality label, we cannot be certain whether the AMNH 
female is from New York or Canada, and so we leave the 
type locality as originally published. We recognize Cato- 


cala residua as a full species. 


Catocala retecta Grote, 1872. Trans. Amer. Entomol. 
Soc. 4:4 


The original description states “A few specimens ex- 
amined from the Middle States.” A male type is at the 
ANSP, and to clarify application of the name we hereby 
designate it as LECTOTYPE for retecta (Fig. 4c). The 
lectotype bears the labels “7”, “TYPE No. 7660/Cato- 
cala/retecta/A. R. Grote”, “LECTOTYPE/Catocala re- 
tecta/Grote 1872/Desig. Gall & Hawks 2002.” The type 
locality is Middle States [USA]. We recognize Catocala 
retecta as a full species. 


Catocala robinsoni Grote, 1872. Trans. Amer. Ento- 

mol. Soc. 4:20 

The original description states “After writing so far my 
kind friend, Mr. Charles A. Blake, sends me specimens 
of two black winged species from Pennsylvania . . . d° 
... There are several old robinsoni at the ANSP, pinned 
in a style like other Grote types at ANSP. A male and fe- 
male in Drawer 3 at the BMNH bear conflicting locality 
and date labels, and no type designations. To clarify ap- 
plication of the name, we hereby designate a male from 
the ANSP collection as LECTOTYPE for robinsoni 
(Fig. 4d). The lectotype bears the labels “Penn.”, “LEC- 
TOTYPE/Catocala robinsoni/Grote 1872/Desig. Gall & 
Hawks 2002.” The type locality is Pennsylvania [USA]. 
We recognize Catocala robinsoni as a full species. 


250 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


IG VI cimens of moths in the genus Catocala Schrank. A. R. Grote types (authorship is Grote unless otherwise noted). a, lecto- 
pon 1 te & Robinson. b, lectotype, residua. e, lectotype, retecta. d, lectotype, robinsoni. e, lectotype, scintillans Grote & Robin- 
son. f, lectotype, seimirelicta. g, lectotype, simulatilis. h, lectotype, sinuosa. 0.9-1.0 x natural size. 


VOLUME 56, NUMBER 4 


Catocala scintillans Grote & Robinson, 1866. Proc. 
Entomol. Soc. Phil. 6:28 


The original description states “Pennsylvania. Coll. 
Ent. Soc. Phil.” In Drawer 3a at the BMNH is a fe- 
male bearing a blue-bordered Grote label, and la- 
beled as type by Butler. This female bears a conflict- 
ing and probably erroneous locality label indicating 
Ohio as the locality. At the ANSP are two old speci- 
mens of scintillans, one female from Ohio ex Strecker 
collection and one unlabeled male pinned in a style 
matching other Grote types at the ANSP. We place 
greater confidence in the blue-bordered label on the 
BMNH female, and hereby designate it as LECO- 
TYPE for scintillans (Fig. 4e). The lectotype bears the 
labels “Ohio”, “Ohio/Grote Coll./81-116.”, “Ohio/81- 
116 [and on the reverse:] Catocala/scintillans/Type 
Grote & Robinson”, “Catocala/scintillans/G + R.”, 
“LECTOTYPE/Catocala scintillans/Grote & Robin- 
son 1866/Desig. Gall & Hawks 2002.” We retain the 
originally published Pennsylvania [USA] as the type 
locality. The name scintillans is a synonym of Catocala 
innubens Guenée, and represents specimens with a 
crisply delimited, wide white distal area on the 
forewing. 


Catocala semirelicta Grote, 1874b. Sixth Ann. Rep. 
Peabody Acad. Sci., p. 35 


The original description does not state the number 
of types. In Drawer 17a at the BMNH is a male bear- 
ing a blue-bordered Grote label and a red-bordered 
type label, and a red BMNH type disc. To clarify ap- 
plication of the name we hereby designate it as LEC- 
TOTYPE for semirelicta (Fig. 4f). The lectotype bears 
the labels “U. S. America/Grote Coll./81-116.”, “This 
spec. was exhibited in the/Can. Coll. at the Centennial 
1876/and has lost its color from exposure”, “United 
States/81-116 [and on the reverse:| Catocala/semire- 
licta/Type Grote”, “Catocala/semirelicta/Grote’”, “Cato- 
cala/semirelicta/Type Grote”, “LECTOTYPE/Catocala 
semirelicta/Grote 1874/Desig. Gall & Hawks 2002.” 
The type locality is Grimsby, Ontario [Canada]. We 
recognize Catocala semirelicta as a full species. 


Catocala simulatilis Grote, 1874c. Trans. Amer. 
Entomol. Soc. 5:94 


The original description states “Ohio, Dr. Hodge, 
two specimens.” In Drawer 2 at the BMNH is a fe- 
male, bearing a red BMNH type disc, and labeled as 
type by Grote. Although this specimen claims to be 
from New York, this is probably a mislabeling, and we 
place greater confidence in Grote’s published state- 
ment. To clarify application of the name we hereby 
designate this specimen as LECTOTYPE for simu- 


251 


latilis (Fig. 4g). The lectotype bears the labels “N. 
Y./Grote Coll./New York’, “New York/81-116 [and on 
the reverse:] Catocala/simulatilis/Grote Type’, “C. 
simulatilis/Type Grote”, “LECTOTYPE/Catocala sim- 
ulatilis/Grote 1874/Desig. Gall & Hawks 2002.” We 
retain the originally published Ohio [USA] as the type 
locality. The name simulatilis is a synonym of Catocala 
obscura Strecker. 


Catocala sinuosa Grote, 1879. Can. Entomol. 11:15 


The original description states “Iwo specimens. 
Florida, Mr. Albert Koebele.” In Drawer 26a at the 
BMNH isa male, bearing a blue-bordered Grote label 
and a red-bordered type label, and a red BMNH type 
disc. To clarify application of the name we hereby des- 
ignate it as LECTOTYPE for sinuosa (Fig. 4h). The 
lectotype bears the labels “U.S. America/Grote 
Coll./81-116.”, “No. 3”, “United States/81-116 [and on 
the reverse:] Catocala/sinuosa/Type Grote”, “Cato- 
cala/sinuosa/Grote”, “Catocala/sinuosa/Type/Grote’, 
“LECTOTYPE/Catocala sinuosa/Grote 1879/Desig. 
Gall & Hawks 2002.” The type locality is Florida 
[USA]. We recognize sinuosa as a subspecies of Cato- 
cala coccinata Grote with lighter forewings and a 
greatly reduced, sometimes obsolete hindwing median 
band. Specimens from peninsular Florida are refer- 
able to sinwosa, whereas nominate coccinata occurs 
elsewhere in North America. We have, however, seen 
occasional specimens similar to sinuosa from the Gulf 
Coast outside of Florida. 


Catocala snowiana Grote, 1876b. Checklist Noct. 
Amer., p.41 


The original description does not state the number 
of types. A female at the AMNH is labeled by Grote as 
type of “snowi,” and to clarify application of the name 
we hereby designate it as LECTOTYPE for snowiana 
(Fig. 5a). The lectotype bears the labels “390”, “No. 
11876/Collection/Hy. Edwards.”, “Type/No./A.M.N.H.”, 
“Catocala/snowi/Type’", “LECTOTYPE/Catocala snow- 
iana/Grote 1876/Desig. Gall & Hawks 2002.” The type 
locality is Kansas [USA]. The lectotype of snowiana is 
a specimen of Catocala palaeogama Guenée. Thus, 
snowiana is hereby transferred from the synonymy of 
Catocala neogama J. E. Smith to that of palaeogama 
(REVISED SYNONYMY). 


Catocala sordida Grote, 1877. Can. Entomol. 9:170 


The original description states “In Mrs. Bridgham’s 
collection is (or rather, was) a specimen labeled similis 
by Mr. Edwards, which belonged to what I consider as 
a variety of gracilis, having the primaries mixed bluish 
bray, and the basal dash of gracilis is wanting. This last 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic.5. Type specimens of moths in the genus Catocala Schrank. A. R. Grote types (authorship is Grote unless otherwise noted). a, lecto- 
type, snowiana. b, lectotype, subnata. ec, lectotype, sordida. d, lectotype, verrilliana. e, lectotype, westcottii. 0.9-1.0 X natural size. 


seems the only important character . . . This dark 
form (which seems also a little shorter winged) has 
been taken with the type by myself near Buffalo, and 
by Dr. Bailey near Albany. I have seen it also from 
Pennsylvania . . . This mixed dark blue-gray form, 
with distinct black lines and without the small basal 
streak of gracilis, I propose to designate by the name 
sordida . A female ex Bailey collection is in 
Drawer 23 at the BMNH, bearing a red-bordered 
type label, and a red BMNH type disc. As Grote cor- 
rectly noted, there is no specimen of sordida labeled 
similis by Edwards in the Bridgham material now at 
the YPM, although there is a series of 12 gracilis. To 
clarify application of the name we designate the fe- 
male at the BMNH as LECTOTYPE for sordida (Fig. 
5c). The lectotype bears the labels “Grote Coll./81- 
IN, “Cena, INNG/fully 12, S77 Dre ||, S. Bate 
ley,/Collector”, “New York/81-116 [and on reverse:] 
C. gracilis/var. sordida/Type Grote”, “var. sordida/ 
Type Grote”, “LECTOTYPE/Catocala sordida/Grote 
1877/Desig. Gall & Hawks 2002.” The type locality is 
hereby restricted to Centre, [Albany County], N[ew] 
Yl[ork, USA] on the basis of the lectotype locality la- 
bel. We recognize Catocala sordida as a full species. 


Catocala subnata Grote, 1864b. Proc. Entomol. Soc. 


Phil 


The original description states “Hab. Maryland. 
(Coll. Ent. Soc. Phil.).” There are two old unlabeled 
males at the ANSP. A female subnata is in Drawer 5 at 
the BMNH, bearing a red BMNH type disc and la- 
beled as type by Butler, but with a conflicting locality 
label “Kansas.” This female also bears a red-bordered 
label without type attribution, stating only “Catocala 


subnata,” and not in Grote’s handwriting. In the same 


series in Drawer 5 is a male subnata bearing a blue- 
bordered Grote label. 

The text of the original description of subnata was 
based on the male, and the accompanying illustration 
in Fig. 4 of Plate 5 is definitely a male, lacking the 
basal dashes and darker scaling on the forewing typical 
of female subnata. The BMNH specimen purporting 
to be a type is a strongly marked female, and it clearly 
does not match the illustration in Fig. 4 of Plate 5. We 
are willing to accept Hampson’s (1913:39) statement 
that the BMNH holds a type of swbnata, but challenge 
his assertion that this is the female with two contradic- 
tory labels, as the male therein bearing Grote’s blue- 
bordered label is a far more plausible candidate. We 
suggest an erroneous association occurred, and we 
hereby designate the male in Drawer 5 as LECTO- 
TYPE for swbnata (Fig. 5b). The lectotype bears the 
labels “U. S. America/Grote Coll./81-116.”, “United 
States/81-116 [and on the reverse:] Catocala/sub- 


VOLUME 56, NUMBER 4 


nata/Grote”, “Catocala/subnata/Grote”, “LECTOTYPE/ 
Catocala sordida/Grote 1877/Desig. Gall & Hawks 
2002.” The type locality is Maryland [USA]. We recog- 
nize Catocala subnata as a full species. 


Catocala verrilliana Grote, 1875a. Can. Entomol. 7:185 


The original description does not state the number 
of types. In Drawer 27 at the BMNH is a male labeled 
as type by Grote, bearing a red BMNH type disc. To 
clarify application of the name we hereby designate it 
as LECTOTYPE for verrilliana (Fig. 5d). The lecto- 
type bears the labels “499”, “U. S. America/Grote 
Coll./81-116”, “United States/81-116 [and on the re- 
verse:] Catocala/verrilliana/Type Grote”, “Catocala/ 
verrilliana/Type/Grote”, “LECTOTYPE/Catocala ver- 
rilliana/Grote 1875/Desig. Gall & Hawks 2002.” The 
type locality is Bosque Co[unty], Texas [USA]. We rec- 
ognize Catocala verrilliana as a full species. 


Catocala westcottii Grote, 1878. Can. Entomol. 
10:195 


The original description discusses two specimens: “I 
have the male from Illinois, Mr. Bean, number 577; the 
female from Wisconsin, Mr. O. S. Westcott, for whom I 
name the species.” The male is in Drawer 25 at the 
BMNH, bearing a red BMNH type disc, and to clarify 
application of the name we hereby designate it as LEC- 
TOTYPE for westcottii (Fig. 5e). The lectotype bears 
the labels “577”, “U. S. America/Grote Coll./81-116.”, 

“United States/81-116 [and on reverse:] Catocala/west- 
cottii/Type Grote”, “Catocala/westcottii/ Grote/Type”, 
“LECTOTYPE/Catocala westcottii/ Grote 1878/Desig. 
Gall & Hawks 2002.” The type locality is hereby re- 
stricted to Illinois [USA] on the basis of the lectotype 
labels. The name westcottii is a synonym of Catocala 
amestris Strecker. 


Infrasubspecific Names 


Catocala habilis var. “basalis” Grote, 1876a. Can. En- 
tomol. 8:230 


The original description states “Specimens received 
from Mr. Robert Bunker, taken about Rochester, N. 
Y.” There is a female in Drawer 5 at the BMNH, la- 
beled as type by Butler and bearing a red BMNH type 
disc, with a locality label “Oneida Co./Aug. 1876/Dr. J. 
S. Bailey,/Collector.” Oneida County is about 80 miles 
from Rochester proper, and is not inconsistent with 
the statement “taken about Rochester.” We leave the 
type locality as Rochester, N[ew] Y[ork USA], as origi- 
nally published. The name “basalis” is a synonym of 
Catocala habilis Grote, and represents the typical fe- 
male of this species. 


Catocala cerogama var. “bunkeri” Grote, 1876a. Can. 
Entomol. 8:230 


The original description does not state the number 
of types. In Drawer 22 at the BMNH is a female bear- 
ing a red-bordered Grote label that lacks the word 
type, but with a type designation by Butler. The type 
locality is New York [USA]. The name “bunkeri” is a 
synonym of Catocala cerogama Guenée, and repre- 
sents specimens with the iano ‘yellow basal shade 
entirely lost.” 


Catocala innubens var. “flavidalis” Grote, 1874c. 
Trans. Amer. Entomol. Soc. 5:95 


The original description states “I have received from 
Prof. S. A. Forbes, Normal, Illinois, a specimen with 
the number ‘2’ attached to it.” In Drawer 3a at the 
BMNH is a male labeled by Grote as “var 
flavidalis/Grote,” and bearing a handwritten label “2.” 
Neither Beutenmiiller (1903a) nor Hampson (1913) 
list the type of “flavidalis” as being among the Nearctic 
Catocala at the BMNH, but Beutenmiiller (1903b) in- 
dicates the type is there, and from the label data we 
are confident the BMNH male is the holotype by 
monotypy. The type locality is not stated in the original 
description, but is presumably [Illinois, USA]. The 
name “flavidalis” is a synonym of Catocala innubens 
Guenée, and represents an aberration with yellowish 
hindwing bands. 


Catocala concumbens ab. “hilli”? Grote, 1883b. 


Papilio 3:43 


The original description states “This very singular 
aberration has been taken by Mr. W. W. Hill at Center, 
N. Y.” This female, the holotype by monotypy, is at the 
New York State Museum (NYSM) in Albany. The type 
locality is Center, [Albany County,] N[ew] Y[ork, 
USA]. The name “hilli” is a synonym of Catocala con- 
cumbens Walker, and represents an aberration with 
yellowish hindwing bands. 


ACHILLE GUENEE 


As was the case with many lepidopteran groups, 
comparative study of the Nearctic species of Catocala 
did not really commence until the 1850's, with the 
publication of A. Guenée’s (1852) Species General and 
F. Walker's ([1858]) List of the specimens of lepi- 
dopterous insects in the collection of the British Mu- 
seum. Guenée described 17 Nearctic Catocala in 
1852, and he and Walker are the most important Eu- 
ropean workers to have published on the Nearctic 
species. Guenée obtained Catocala for his own collec- 
tion from several North American sources, and had 


the opportunity to study Boisduval’s collection and that 
of the BMNH, which included Doubleday and Abbot 
material. Most of Guenée’s original descriptions cite 
specimens, but he also described two Nearctic Cato- 
cala from unpublished paintings by John Abbot. 

Identifying many of Guenée’s noctuid species was 
an odyssey for Nearctic workers. Although Grote trav- 
eled to France to visit Guenée and examine his type 
material, Grote was unable to resolve noctuids that 
Guenée had based on Abbot paintings. In a biography 
of Guenée, he summarized his opinion as follows 
(Grote 1881b): “[Guenée] drew up descriptions of sev- 
eral species [of North American noctuids] from draw- 
ings by Abbot, and none of these have been satisfacto- 
rily identified and probably cannot be.” J. B. Smith 
toured many of the European collections for his noc- 
tuid catalogue, and stated therein (1893:9) “I did suc- 
ceed in getting at most of the species marked ‘M.N., 
in the Species General. I was in hopes of being able to 
get track of the Abbot drawings, from which Guenée 
described a number of species; but I did not succeed 
in this;” he also indicated that while at the BMNH his 
“time was limited, I did not study Acronycta, which 
Dr. Riley had already studied on a prior occasion, nor 
Catocala, which Messrs. Grote and Henry Edwards 
had compared.” 

The first person to claim to know the identity of the 
Guenée Catocala names connubialis, messalina and 
micronympha was G. D. Hulst (1884). Hulst was unfa- 
miliar with these taxa just a few years earlier (Hulst 
1880), and apparently based his 1884 opinions not on 
comparisons of specimens but on a letter he received 
in the intervening years from A. G. Butler of the 
BMNH. Hulst’s prior nomenclatural actions in the 
genus had already come under blistering attack by 
Grote (1881c), and Hulst’s rather sudden “understand- 
ing” of the Guenée names simply became additional 
fodder for the long-running feud between these two 
workers (e. (ee Grae 1891, 1892: Hulst 1881, 18932). 
Nevertheless, over the ensuing several decades, 
Hulst’s concepts of connubialis, messalina, and mi- 
cronympha became fixed in the Nearctic literature. 

In - Cae of messalina, Hulst (1884:34) stated 
that “Mr. Charles Oberthur, of Rennes, France, to 
ee Cnencek types came, informs me that when 
they came into his possession, all were found to be de- 
stroyed beyond the possibility of recovery by the larvae 
of Anthrenus |dermestids]. So no identification can be 
made of any of Guenée’s American species described 
from imagines, from the types themselves.” Although 
Hulst’s statement may be true for messalina, since we 

e not located any types, it is demonstrably false for 


Guenée’s other Catocala, as W. Barnes obtained both 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


these and Guenée’s other North American moth types 
from C. Hofer in 1927 prior to the transfer of the 
Oberthur collection to the BMNH. The Guenée Cato- 
cala types are at the USNM, bearing Guenée, Oberthur, 
and Barnes collection/accession labels, as well as 
Guenée’s distinctive handwritten type labels in French. 

Because Guenée described a number of lepi- 
dopteran species based on paintings by John Abbot, 
we wish to elaborate here on the pioneering work of 
this man, one of the most productive early naturalists 
in North America during the late 18th and early 19th 
centuries. John Abbot reared, collected, and painted 
native birds, insects, and plants of the southeastern 
United States, and was one of the primary sources of 
specimen material for customers in Europe and North 
America at the time. He executed thousands of paint- 
ings, the vast majority of which was never published. A 
meticulous and revealing book on Abbot's life and ac- 
complishments was published recently by Gilbert 
(1998). 

Only four of Abbot’s paintings of Nearctic Catocala 
were published, in The Natural History of the Rarer 
Lepidopterous Insects of Georgia (Smith [& Abbot] 
1797). In that tome, four new species of Catocala were 
described from the four Abbot paintings, these being 
amasia (J. E. Smith), consors (J. E. Smith), neogama, 
and vidua (J. E. Smith). The identity of the latter three 
names as separate species was settled quickly by the 
19th century Nearctic Catocala workers, whereas the 
applicability of the name amasia was not completely 
resolved until well into the 20th century (see Gall 
1992, and our treatment below of connubialis Guenée; 
the name amasia J. E. Smith was recently suppressed 
[ICZN 1994]). 

Abbot painted many other species of Nearctic Cato- 
cala, and these paintings survive today in several loca- 
tions. The two most notable compilations are the 
bound volumes in the Entomology Library of the 
BMNH (ex Francillon collection) and the Houghton 
Library at Harvard University (ex Oemler collection), 
both of which we have examined. Among the 17 
quarto volumes at the BMNH are 14 plates illustrating 

18 individual Catocala, and at Houghton are 16 plates 
illustrating 21 individual Catocala (see Table 2). To- 
gether, the BMNH and Houghton plates depict at 
east 18 different species. Ten plates are common to 
both BMNH and Houghton, showing the same species 
on the same backgrounds, and these likely represent 
standard illustrations that Abbot's clients could request 
(see Gilbert 1998). 

Perhaps not surprisingly, given his keen eye and 
long-term residency in Georgia, Abbot knew of and 
painted several rare and/or local Catocala species from 


VOLUME 56, NUMBER 4 


255 


Fic. 6. Type specimens of moths in the genus Catocala Schrank. A. Guenée types. a, lectotype, andromedae. b, lectotype, androphila. e, 
neotype, connubialis. d, lectotype, cerogama. e, lectotype, desperata. £, lectotype, melanympha. g, neotype, micronympha. h, holotype, 
polygama. i, lectotype, palaeogama. j, lectotype, parta. 0.9-1.0 x natural size. 


the southeastern United States that were not “discov- 
ered” and described until late in the 20th century. For 
example, one of the plates present at both BMNH and 
Houghton clearly shows Catocala grisatra Brower (un- 
described until 1936) at the top and Catocala similis at 
the bottom. Among the non-corresponding plates, one 
at Houghton depicts Catocala louiseae Bauer (unde- 


scribed until 1965), and another at Houghton appears 
to depict the as yet undescribed species in the Cato- 
cala amica complex. At this juncture, we believe none 
of the 19th and early 20th century Nearctic Catocala 
workers studied Abbot’s unpublished paintings; 
species such as grisatra and louiseae are, in our opin- 
ion, far too distinctive to have escaped being named, 


256 


TABLE 2. The identity of Nearctic Catocala Schrank species fig- 
ured in unpublished paintings by John Abbot, in the bound volumes 
housed at the Houghton Library (Harvard University, Cambridge, 
Massachusetts) anh the Entomology Library at the Natural History 
Museum (London, England). For plates with two listed names, the 
first appears above the second on the plate. 


Source Taxon 


Houghton Library (Oemler) 


pl. 123 epione (Drury) 

pl. 124 ilia (Cramer) 

pl. 125 ultronia (Hiibner) 

pl. 126 consors J. E. Smith, muliercula Guenée 
pl. 127 louiseae Bauer 

pl. 128 lineella? Grote, jair? Strecker 

pl. 142 sappho Strecker, maestosa Hulst 

pl. 143 andromedae Guenée 

pl. 144 amatrix (Hiibner) 

pl. 145 carissima Hulst 

pl. 146 coccinata Grote 

pl. 147 nebulosa W. H. Edwards 

pl. 148 consors J. E. Smith, muliercula Guenée 
pl. 149 grisatra Brower, similis W. H. Edwards 
pl. 150 micronympha Guenée 

pl. 151 amica (Hiibner), jair? Strecker 


Natural History Museum (Francillon) 
vol, 8, pl. +52 carissima Hulst 
vol. 8, pl. 264 amatrix (Hiibner) 


vol. 8, pl. 265 coccinata Grote 

vol. 8, pl. 266 maestosa Hulst 

vol. 8, pl. 267 nebulosa W. H. Edwards 
vol. 8, pl. 270 amica (Hiibner) 

vol, 8, pl. 271 micronympha Guenée 
vol, 8, pl. 273 andromedae Guenée 


a 
vol. 17, pl. 29 
vol. 17, pl. 30 
vol. 17, pl. 89 
vol. 17, pl. 126 
vol. 17, pl. 140 
vol. 17, pl. 280 
vol. “Tring”, pl. 23 
vol. “Tring”, pl. 24 


carissima Hulst 

sappho Strecker, maestosa Hulst 

ilia (Cramer), ilia (Cramer) 

amica (Hiibner), amica (Hiibner) 
grisatra Brower, similis W. H. Edwards 
consors J. E. Smith, muliercula Guenée 
amatrix (Hiibner) 

coccinata Grote 


had the paintings been seen. One complicating aspect 
of the bound Francillon and Oemler volumes is that 
the plates are labeled haphazardly. For example, at the 
BMNH are plates bearing the labels vidua (Plate 30), 
amasia (Plate 126), consors (Plate 140) and neogama 
(Plate 280), that is, the four J. E. Smith names of 1797. 
However, these plate names do not in any way match 
Smith's published usages. Instead, Plate 30 depicts 
sappho Strecker, Plate 126 depicts amica, Plate 140 
depicts grisatra and similis and Plate 280 depicts con- 
sors and muliercula Guenée. In addition, plates often 
bear confusing subsequent annotations. For example, 
at Houghton, “C. marmorata>?” appears on Plate 123 
next to a painting of Catocala ilia (Cramer), and 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


“grynea? Cr.” appears on Plate 127 next to a painting 
of Catocala louiseae. 

There are two Nearctic Catocala names whose de- 
scriptions were made from reference to unpublished 
Abbot paintings. Both names were authored by 
Guenée (1852), these being Hypogramma androme- 
dae and Catocala connubialis. Hulst (1884) first placed 
the name connubialis into the synonymy of Nearctic 
Catocala under his new species sancta Hulst, and 
Barnes and McDunnough (1918b) later introduced 
andromedae as the senior synonym of tristis Edwards, 
based on information from Hampson. Guenée’s origi- 
nal description of andromedae is distinctive, and 
Hampson’s association of this name with the previ- 
ously published name tristis has been universally ac- 
cepted. We have located unpublished Abbot plates 
matching the description of andromedae. In contrast, 
Guenée’s original description of connubialis could ap- 
ply to infrapopulational morphs of at least two Cato- 
cala species, and Hulst’s association of this name with 
the previously published amasia and cordelia Hy. Ed- 
wards met with immediate resistance. Fortunately, 
Hulst’s concept of connubialis is readily established 
from his type of sancta. However, we have not located 
an Abbot painting that matches Hulst’s concept. We 
discuss these situations below. 

We note here that the Entomology Library at the 
BMNH contains a bound volume, in excess of 450 
pages and handwritten by Guenée in French, that con- 
tains extensive notes by him on the synonymy of hol- 
arctic Lepidoptera (Guenée [ca. 1857]). It also contains 
descriptions with manuscript names for non-EKuropean 
species that were never published, including Catocala. 
We were unable to glean any additional data about 
Guenée’s published Nearctic Catocala names from this 
volume, but we recommend that lepidopteran system- 
atists routinely consult this work when studying 
Guenée types, as it clearly holds significant informa- 
tion that complements the Species General. 


Available Names 


Catocala andromedae Guenée, 1852. Hist. Nat. Spec. 
Gen. Lepid. 7:36 


Guenée described Hypogramma andromedae from 
a painting by John Abbot. The name andromedae was 
not included in the major works on Nearctic Catocala 
during the 19th century, probably because the species 
was originally described in another genus. The combi- 
nation Catocala andromedae appears to have been 
used first in Barnes and McDunnough’s (1917) check- 
list, and the following year in their Catocala mono- 


VOLUME 56, NUMBER 4 


graph, where they stated (1918b:36) “This species has 
been generally known as tristis Edwards but Sir Geo. 
Hampson has recently called our attention to the fact 
that Guenée’s description of andromedae (a name 
heretofore unplaced) fits this species very well; we 
concur with him in using Guenée’s name, which has 
priority.” We agree with this placement, and further 
note that the larval illustration of andromedae by 
Guenée (1852, fig. 11, Plate 2) is a Catocala and 
clearly matches larvae in the small group of Nearctic 
Ericaceae-feeding species, to which tristis belongs. 
We have recently examined the compilations of Ab- 
bot’s unpublished works at the BMNH and the 
Houghton Library at Harvard University. Paintings 
that are readily identifiable as tristis appear in both 
compilations. To clarify application of the name we 
hereby designate the specimen figured (and now pre- 
sumed lost) on Plate 273 in bound Abbot Volume 8 at 
the BMNH as LECTOTYPE for andromedae (Fig. 
6a). The accompanying text reads “No 273. Noctua. 
Numeria. Caterpillar feeds on Oaks, bred 10th June, 
frequents Hammock, Rare, called in Savannah, small 
black underwing.” The foodplant association is erro- 
neous, as are a number of Abbot’s. Guenée’s original 
description states “Amerique Septentrionale,” and on 
the basis of the text for Plate 273 and the published ac- 
counts of Abbot’s activities (see Gilbert 1998) we 
hereby restrict the type locality to Georgia, USA. We 
recognize Catocala andromedae as a full species. 


Catocala androphila Guenée, 1852. Hist. Nat. Spec. 
Gen. Lepid. 7:106 


The original description states “Amerique Septen- 
trionale. Coll. Gn.” Three Guenée males labeled as 
androphila are at the USNM. Two are Catocala am- 
ica, and one is Catocala lineella. Guenée’s large label 
in French is affixed to an amica, and indicates that 
the first two specimens in his series (both amica) are 
“ordinaire” whereas the third in the series (the li- 
neella) is “var A. Gn. p. 107” (this confirms that Grote 
1872:18 had correctly surmised that his new species 
lineella was Guenée’s var A). The label also says 
“Amer. Boreale. 1845,” and then “I have not seen 
since then a large enough number, all alike, sender 
also [unintelligible] Baltimore, but I have not seen 
the female.” To clarify application of the name we 
hereby designate the male bearing Guenée’s large la- 
bel in French as LECTOTYPE for androphila (Fig. 
6b). In addition to Guenée’s label, the lectotype bears 
the labels “Ex Musaeo/Ach. Guenée’”, “Oberthur/Col- 
lection”, “Barnes/Collection”, “LECTOTYPE/Cato- 
cala androphila/Guenée 1852/Desig. Gall & Hawks 


2002.” The type locality is hereby amended to 
Amer[ique] Boreale on the basis of the lectotype local- 
ity label. Barnes and McDunnough (1918b:45) 
pointed out that Guenée intended the name an- 
drophila to replace the name amica (Hiibner), which 
Guenée mistakenly believed was preoccupied by the 
Palearctic species Hadena amica Treitschke (1825). 
The name androphila is a synonym of and unnecessary 
replacement name for Catocala amica (Hiibner). 


Catocala cara Guenée, 1852. Hist. Nat. Spec. Gen. 

Lepid. 7:87 

The original description states “Coll. Gn. Un d.” A 
male bearing a large Guenée label in French stating 
“No. 1” is at the USNM. This specimen also bears an- 
other label “Type/see label” by F. H. Benjamin, and we 
consider it to be the holotype by monotypy. The type 
locality is the vicinity of Baltimore [Maryland, USA]. 
We recognize Catocala cara as a full species. 


Catocala cerogama Guenée, 1852. Hist. Nat. Spec. 
Gen. Lepid. 7:96 


The original description states “Coll. Feisth. Gn. et 
M. N.” A male and female are at the USNM, both 
bearing Guenée labels in French. The two labels were 
once part of a single label, and indicate: “[on the male] 
1. Poor d from the collection of Feisthamel. / [on the 
female] 2. Good ° vicinity of Baltimore, purchased M. 
Becker in 1852.” To clarify application of the name we 
hereby designate the female as LECTOTYPE for 
cerogama (Fig. 6d). In addition to Guenée’s label, the 
lectotype bears the labels “Ex Musaeo/Ach. Guenée’, 
“Oberthur/Collection”, “Barnes/Collection”, “LEC- 
TOTYPE/Catocala cerogama/Guenée  1852/Desig. 
Gall & Hawks 2002.” The type locality is hereby re- 
stricted to the vicinity of Baltimore [Maryland, USA] 
on the basis of the lectotype locality label. We recog- 
nize Catocala cerogama as a full species. 


Catocala connubialis Guenée, 1852. Hist. Nat. Spec. 
Gen. Lepid. 7:105 


Guenée described Catocala connubialis from a 
painting by John Abbot. Eventually, nearly a century 
later, as a result of life history and related studies, the 
names amasia and connubialis were recognized to be 
infrasubspecific morphs (along with cordelia, sancta 
and virens French) of one of the most variable Nearc- 
tic Catocala species. Since 1938, the name connubialis 
has been applied to the species because amasia (pub- 
lished by J. E. Smith in 1797) was considered preoccu- 
pied by Catocala amasia Esper (1787), an unrelated 
Palearctic species thought to have been published in 


1796. However, Heppner (1981) showed that Esper's 
work dates from 1804, not 1796. The applicable his- 
tory was reviewed by Gall (1992) in a petition accepted 
by the ICZN (1994) to conserve the name connubialis 
Guenée and suppress the name amasia J. E. Smith. 

As noted above, Hulst was the first Nearctic Cato- 
cala worker to claim to understand the identity of con- 
nubialis. He first mentioned the name (1884:34) in his 
account of Catocala messalina: “Connubialis hereto- 
fore not identified, was described from a colored draw- 
ing of Abbott which is now, I understand, in the British 
Museum.” Later in the same paper, Hulst (1884:3S) 
described Catocala sancta as new, placing connubialis 
as its synonym. The description of sancta follows: 
“These last two species |amasia and similis | have been 
much confused. Abbott’s upper figure, from which 
Smith says his description was anagle. is undoubtedely 
the insect afterwards named cordelia by Hy. Edwar ds. 
Guenée describes the lower figure of Abbott as ama- 
sia; but Abbott's description of the upper figure, holds 
good against that of Guenée. Mr. Grote identified the 
more southern form as amasia, and thus it is generally 
labelled in collections. Mr. A. G. Butler [of the 
BMNH] writes me, this latter is connubialis, Guen.: 
but the description does not fit, and it was described 
from a drawing, and so the name does not in any case 
hold.” At the end of the same paper, Hulst (1 884:56) 
commented further on the Butler letter, stating “Page 
38, line 15, after ‘writes me’ insert ‘his opinion is.’ I do 
not understand that he has ultimate evidence. At any 
rate the description being based on a picture, can not 
stand.” 

Hulst (1892:74) later elaborated his position: “Prof. 
French says ‘var. Virens is not a variety of Cordelia, Hy. 
Edw., but of Amasia; and Cordelia is not the one fig- 
ured by Dr. Strecker, pl. 9, f. 12.” But cordelia, Hy. 
Edw., is a synonym of amasia, Ab. & Sm., and Dr. 
Strecker’s figure is not amasia, Ab. & Sm. The error 
comes from the fact that Abbot & Smith figured two 
species as male and female of amasia, the description 
being of the upper one only. The insect represented by 

the lloxer figure of Abbot & Smith was distributed by 
Mr. Grote, and figured by Dr. Strecker as amasia. Of 
course the name attaches to the figure described, as af- 
terwards Guenée located it, calling the lower figure 
connubialis. The lower insect I afterwards described as 
sancta, regarding Guenée’s name as without authority, 
as the description was from a picture. Whether I was 
right or not I will not here say, but the insect distrib- 
uted by Mr. Grote, and figured by Dr. Strecker as ama- 

ia, is either connubialis, Gn., or sancta, Hulst: while 
Abbot & Smith is the cordelia of Hy. Ed- 
wards, as Mr. Edwards afterwards acknowledged to 


the amasia oi 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


me.” This last sentence is Hulst’s most succinct diag- 
nosis of connubialis. 

The two species shown on Smith and Abbot's Plate xe 
are indeed not conspecific, and not even that close mor- 
phologically. This was first noted by Grote and Robin- 
son (1866), who restricted the name amasia to the up- 
per illustration and applied the name formula to the 
lower illustration (formula was later transferred to the 
synonymy of similis). This interpretation of Plate xc 
was followed by all the Nearctic Catocala workers, and 
Hulst (1884:37) had already unequivocally adopted 
this interpretation in his synonymies for both similis 
(“pl. 90 lower figure”) and amasia (“pl. 90, upper fig.”). 
Hulst described sancta in the same 1884 paper as dis- 
tinct from both similis and amasia, and his type of 
sancta is identical to Strecker’s Plate 9 Fig. 12 and not 
in any way confusable with similis. Yet later, Hulst 
(1892:74) equally unequivocally assigned his sancta to 
the lower illustration of Plate xc. Unless one is willing 
to assume that by 1892 Hulst felt his sancta was con- 
specific with similis, for which we have no evidence 
whatsoever, his 1854 and 1892 synonymic treatments 
appear to be irreconcilable. 

In his noctuid catalogue, Smith (1893:334) did not 
resolve this problem when he indicated: “Dr. Hulst 
cites this species [connubialis], as well as amasia} Grt., 
to sancta, perhaps not justly. There is really no evi- 
dence that Guenée intended the amasiat, and indeed, 
Dr. Hulst says the description does not fit. It [connu- 
bialis | should not therefore be cited as a synonym to 
sancta, which it must othenvise replace. Guenée’s 
species based on Abbot's figures have been universally 
accepted, and if the figures are still in existence there 
is no reason why positive knowledge should not yet be 
obtainable.” 

Even though Guenée’s concept of connubialis had 
not yet been adequately established, Smith’s statement 
probably cemented the link between connubialis and 
sancta. Hulst’s contemporaries could readily deter- 
mine Hulst’s concept of connubialis by reference to 
the type of sancta and Strecker’s illustration on his Fig- 
ure 12 on Plate 9. On the other hand, Hulst’s own un- 
derstanding of connubialis was apparently based solely 
on the letter sent to him by Butler, and this letter was 
undoubtedly not generally available to other Catocala 
workers at the time (we have been unable to locate any 
letters from Butler to Hulst in the archives of Rutgers 
College, or in the Butler correspondence at the 
BMNH). Moreover, Butler’s concept of connubialis, 
and hence Hulst’s, unquestionably stems from 
Walker's List (1858:1207-1208), in which specimen 
material at the BMNH is attributed to connubialis. In- 
deed, in Drawer 32a at the BMNH are two similar 


VOLUME 56, NUMBER 4 


males labeled as connubialis by Butler. These BMNH 
specimens match Hulst’s type of sancta at the AMNH 
and Strecker’s illustration, and are also consistent with 
Guenée’s original description of connubialis. The un- 
dersurface of Butler’s label on one of the males reads 
“Catocala/connubialis var./Type Walker.” 

We had hoped to locate an Abbot painting that 
matched the BMNH specimens Butler labeled as con- 
nubialis, but we found no matching painting at either 
the BMNH or at the Houghton Library, nor in the col- 
lections of Abbot Catocala plates at the CMNH, the 
ANSP, or the Atlanta History Center. Because (a) no 
Abbot painting unequivocally attributable to connu- 
bialis has been located, (b) the original description of 
connubialis is also consistent with some infrapopula- 
tional morphs of Guenée’s micronympha, (c) no type 
for micronympha has been located, and (d) the early 
Nearctic Catocala workers had trouble differentiating 
the small yellow-hindwinged species, we consider it 
essential to fix the name connubialis firmly. To clarify 
application of the name, we hereby designate the 
BMNH male labeled as type of the variety by Butler as 
NEOTYPE for connubialis (Fig. 6c). The neotype 
bears the labels “United States/-? [and on the reverse: | 
Catocala/connubialis var./Type Walker”, “NEO- 
TYPE/Catocala connubialis/Guenée 1852/Desig. Gall 
& Hawks 2002.” Guenée gives a type locality of 
“Amerique septentrionale,” but because Abbot does 
not seem to have ever figured this morph, and the neo- 
type bears no precise locality data, we leave the type 
locality as stated by Guenée. We recognize Catocala 
connubialis as a full species. 


Catocala desperata Guenée, 1852. Hist. Nat. Spec. 
Gen. Lepid. 7:95 


The original description states “Amerique Septen- 
trionale, environs de Baltimore. Coll. Gn. Un ¢, une 
2.” A Guenée male labeled “desperata/Baltimore” is at 
the USNM, and to clarify application of the name we 
hereby designate it as LECTOTYPE for desperata 
(Fig. 6e). In addition to Guenée’s label, the lectotype 
bears the labels “Ex Musaeo/Ach. Guenée’, 
“Oberthur/Collection”, “Barnes/Collection”, “LEC- 
TOTYPE/Catocala desperata/Guenée 1852/Desig. 
Gall & Hawks 2002.” The type locality is the vicinity of 
Baltimore [Maryland, USA]. The name desperata is a 
synonym of Catocala vidua J. E. Smith. 


Catocala innubens Guenée, 1852. Hist. Nat. Spec. 
Gen. Lepid. 7:95 


The original description states “Amerique Septen- 
trionale. Coll. Bdv. Un d. Parait rare.” A male labeled 
“Inmnubens/Gn. 25” and “Ex Musaeo/A. Kuwert 1894” 


is at the USNM (Oberthur obtained the noctuid por- 
tions of Kuwert’s [1828-1894] collection, and other 
Guenée Catocala type material at the USNM bears 
Kuwert collection labels). We consider this USNM 
male to be Guenée’s holotype by monotypy, since the 
handwriting on the first label matches that on other 
USNM Guenée types. The type locality is “Amerique 
Septentrionale.” We recognize Catocala innubens as a 
full species. 


Catocala insolabilis Guenée, 1852. Hist. Nat. Spec. 
Gen. Lepid. 7:94 


The original description states “Coll. Gn. Une ®... 
[Variety] A... Coll. Gn. Un 4,” indicating a female 
holotype by monotypy. There are four male insolabilis 
at the USNM that are apparently from Guenée’s col- 
lection, although only two bear his accession labels. 
We have not located a female, and Guenée appears to 
have accurately sexed all his other Nearctic Catocala 
types. The first Guenée male in the USNM series is 
worn, and the second male is in good condition. The 
worn male bears a large Guenée label in French that 
states: “Baltimore 2 M. Becker. The ¢ that I have de- 
scribed is [unintelligible] poor [unintelligible] into a 
collection. I doubt [unintelligible] variety, in all, as I 
have supposed, [unintelligible] the No. 1 ¢ which ar- 
rives since is quite similar to the female.” We believe 
the worn male is the one on which Guenée’s variety A 
was based. In the original description, Guenée was un- 
certain whether variety A represented the typical male 
of the species, but his specimen label seems to corrob- 
orate a correct association of the sexes. Because the 
original description is diagnostic, and there has been 
no confusion regarding the applicability of the name, 
we choose not to take any formal action. The type lo- 
cality is hereby restricted to Baltimore [Maryland, 
USA] on the basis of Guenée’s label. We recognize 
Catocala insolabilis as a full species. 


Catocala lacrymosa Guenée, 1852. Hist. Nat. Spec. 
Gen. Lepid. 7:93 


The original description states “Coll. Gn. Un beau 
d.” This Guenée male, the holotype by monotypy, is at 
he USNM. Its large Guenée label in French offers no 
substantive accastonell information beyond that pro- 
vided in the original description. The type locality is 
“Amerique Septentrionale.” We recognize Catocala 
lacrymosa as a full species. 


Catocala melanympha Guenée, 1852. Hist. Nat. 
Spec. Gen. Lepid. 7:98 


The original description states “Canada. Coll. Gn. 
Parait rare.” Three Guenée males labeled melanympha 


are at the USNM, and to clarify application of the name 
we hereby designate the male bearing Guenée’s large 
label in French as LECTOTYPE for melanympha (Fig. 
6f). In addition to Guenée’s label, which indicates the 
specimen was sent by Feisthamel, the lectotype bears 
the labels “Ex Musaeo/Ach. Guenée”, “Oberthur/Col- 
lection”, “Barnes/Collection”, “LECTOTYPE/Catocala 
melanympha/Guenée 1852/Desig. Gall & Hawks 2002.” 
The type locality is Canada. The name melanympha is a 
synonym of Catocala antinympha (Hiibner). 


Catocala messalina Guenée, 1852. Hist. Nat. Spec. 
Gen. Lepid. 7:107 


The original description states “Amerique Septen- 
trionale. Coll. Bdv. Un d.” Grote (1872:19) translated 
Guenée’s original description of messalina, and stated 
“Not identified by me. . . . This species should be 
recognisable from the continuous hind border of the 
secondaries, an exceptional character of this group.” 
Hulst (1884) embellished Grote’s translation but cor- 
rectly identified the species, listing jocasta Strecker 
and belfragiana Harvey as synonyms. We have been 
unable to locate a type of messalina at the USNM or at 
the BMNH. However, the original description of this 
unique species is diagnostic, and there has been no 
confusion regarding the applicability of the name since 
Hulst's diagnosis, so we choose not to take any formal 
action. The type locality is “Amerique Septentrionale.” 
We recognize Catocala messalina as a full species. 


Catocala micronympha Guenée, 1852. Hist. Nat. 
Spec. Gen. Lepid. 7:102 


As with connubialis and messalina, Hulst (1884:34) 
was the first Nearctic worker to claim to know the 
identity of micronympha. However, unlike connubialis 
and messalina, Hulst offered no justification whatever 
for his placement of micronympha, stating only “An 
extraordinarily variable species. Atarah is slightly 
lighter than type form.” Although Grote (1891:281) 
properly protested that “without the slightest reason, 
Mr. Hulst quotes fratercula [the previous oldest name] 
as the species intended by Guenée as micronympha,” 
the name micronympha was used in the catalogue by 
Smith (1893) and as the species name by subsequent 
authors. 

The original description of micronympha states 
“Amerique Septentrionale. Un d.” There are no 
Guenée specimens labeled by Guenée as mi- 
cronympha among the type material at either the 
USNM or the BMNH. However, there are two worn 
male Guenée specimens of micr onympha at the 
USNM that are labeled by Guenée as amasia, and the 
large label in French on one male indicates it was used 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


for his 1852 description of amasia. The confusion 
among the 19th century Catocala workers regarding 
amasia sensu Guenée (which does not correspond well 
to Abbot's painting of amasia J. E. Smith) is now much 
more readily understandable, given the label data affil- 
iated with the two USNM micronympha. These two 
USNM micronympha represent an infrapopulational 
variant lighter than “hero” Hulst that occurs in mi- 
cronympha from the southern United States viz., 
forewings of a dull cream color, with variable brown 
shading in the postmedial area and the outer margin. 

Because (a) we have not located the micronympha 
holotype, (b) Hulst’s placement of micronympha was 
made without substantiating published evidence, (c) 
the original description of micronympha is consistent 
with some infrapopulational morphs of connubialis, 
and (d) the early Nearctic Catocala workers had trou- 
ble differentiating the small yellow-hindwinged 
species, we consider it essential to fix the name mi- 
cronympha firmly. Accordingly, we hereby designate a 
male from the AMNH as NEOTYPE for mi- 
cronympha (Fig. 6g). The neotype bears the labels 
“USA: Georgia: Liberty Co./St. Catherines Island/May 
1991/Rozen, Quinter & Sharkov”, “NEOTYPE/Cato- 
cala micronympha/Guenée 1852/Desig. Gall & Hawks 
2002.” The type locality is hereby amended to S[ain]t 
Catherines Island, Liberty Co[unty], Georgia, USA on 
the basis of the neotype label. We recognize Catocala 
micronympha as a full species. 


Catocala muliercula Guenée, 1852. Hist. Nat. Spec. 

Gen. Lepid. 7:97 

The original description states “Amerique Septen- 
trionale. C ‘oll. Bdv. Un d.” We have not been able to lo- 
cate Guenée’s holotype at either the USNM or the 
BMNH. However, the original description is diagnos- 
tic, and we further note that the larval illustration of 
muliercula by Guenée (1852, fig. 15, Plate 2) matches 
the distinctive larvae in the small Nearctic group of 
Myricaceae-feeding species, to which muliercula be- 
longs. Since there is no indication in the early Nearctic 
Catocala literature of confusion as to the applicability 
of this name, we choose not to take any formal action. 
The type locality is “Amerique Septentrionale.” We 
recognize Catocala muliercula as a full species. 


Catocala palaeogama Guenée, 1852. Hist. Nat. Spec. 
Gen. Lepid. 7:97 


The original description states “Amerique Septen- 
trionale. Coll. Bdv. et Feisth.” and then “A ... Memes 
localities. Coll. Gn. Un d.” (Guenée’s variety A is pha- 
langa Grote). There are two male Guenée palaeogama 
at the USNM. One male bears a large Guenée label in 


VOLUME 56, NUMBER 4 


French that discusses four specimens, and indicates “d 
vicinity of Baltimore purchased Becker;” the other 
male is labeled simply “palaeogama.” To clarify appli- 
cation of the name we hereby designate the male bear- 
ing Guenée’s large label as LECTOTYPE for 
palaeogama (Fig. 6i). In addition to Guenée’s label, 
the lectotype bears the labels “Ex Musaeo/Ach. 
Guenée’”, “Oberthur/Collection”, “Barnes/Collection”, 
“LECTOTYPE/Catocala palaeogama/Guenée/Desig. 
Gall & Hawks 2002.” The type locality is hereby re- 
stricted to the vicinity of Baltimore [Maryland, USA] 
on the basis of the lectotype locality label. We recog- 
nize Catocala palaeogama as a full species. 


Catocala parta Guenée, 1852. Hist. Nat. Spec. Gen. 

Lepid. 7:84 

The original description states “Amerique Septen- 
trionale, Canada. Coll. Gn. Trois examplaires.” Three 
male and one female Guenée parta are at the USNM. 
Two males and one female bear sections of what had 
previously been a single large Guenée label in French, 
and the fourth male only bears a label stating 
“Canada.” The three Guenée label segments together 
indicate “Amerique du Nord. 1. male purchased from 
Becker? / 2. 2 id. id. in 1851. Baltimore. / 3. 6 poor 
from coll. Feisthamel.” To clarify application of the 
name we hereby designate the female as LECTO- 
TYPE for parta (Fig. 6j). In addition to Guenée’s label, 
the lectotype bears the labels “Ex Musaeo/Ach. 
Guenée’”, “Oberthur/Collection”, “Barnes/Collection’ , 
“LECTOTYPE/Catocala parta/Guenée 1852/Desig. 
Gall & Hawks 2002.” The type locality is hereby re- 
stricted to Baltimore [Maryland, USA] on the basis of 
the lectotype locality label. We recognize Catocala 
parta as a full species. 


Catocala polygama Guenée, 1852. Hist. Nat. Spec. 
Gen. Lepid. 7:105 


The original description states “Amerique Septen- 
trionale, Canada. Coll. Bdv. Un d.” We have not been 
able to locate Guenée’s type at either the USNM or 
the BMNH. Although polygama has been tabulated as 
a junior synonym of Catocala grynea (Cramer 1780) 
since the early part of the 20th century, reanalysis of 
the original description and especially of Guenée’s ac- 
companying illustration on Fig. 2 of Plate 16 (repro- 
duced in our Fig. 6h) reveals that polygama is in fact 
an unused senior synonym of Catocala alabamae 
Grote (1876a). A petition is currently pending at the 
ICZN (Gall 2002) to suppress the name polygama and 
conserve the name alabamae. We review the history 
and evidence here. 

Following Guenée’s description, Grote (1872:15-16) 


applied polygama to “[specimens from] (Canada to Vir- 
ginia) which differ in appearance among themselves 
but which I cannot separate . . . I think we have to do 
with a single variable species.” Grote distributed speci- 
mens determined by him as polygama to many work- 
ers, and thus the name polygama sensu Grote became 
widely used for a common and well-collected Catocala 
species from northeastern North America—viz, what 
lepidopterists currently call blandula Hulst. For ex- 
ample, Saunders (1876:72) described Catocala crataegi 
from Ontario, Canada as a new species, and provided 
detailed larval and adult descriptions of both it and 
polygama, showing significant points of distinction; 
Saunders’ description of the larva of polygama is a pre- 
cise match for the larva of blandula, and matches the 
larva of no other Catocala species from eastern 
Canada. 

In the first of his two treatises on Catocala, Hulst 
(1880:6-7) placed Grote’s alabamae as a variety of 
grynea; and pretiosa, crataegi and mira Grote (1876a) 
as “sub-varieties” of polygama. In his second treatise, 
Hulst (1884:35-39) returned crataegi and mira to full 
species status, and noted, correctly, that Grote had 
misidentified Guenée’s polygama. To resolve this, 
Hulst placed polygama sensu Guenée as a synonym of 
grynea, and proposed the new name blandula for the 
previously misidentified species polygama sensu 
Grote. Hulst (1884:35) did not unequivocally resolve 
polygama sensu Guenée, as indicated by his use of “C. 
polygama, Guen. Noct. 7, 105, pl. 16, f.2, (?)” in his 
synonymy for grynea. Hulst (1884:36) elaborated: 
“The description of polygama, Guen., seems to fit this 
species [grynea]; the figure [given by Guenée], which 
is poor, seems more like var. alabamae; neither de- 
scription nor figure approach the insect identified as 
polygama by Grote.” Under his treatment of blandula, 
Hulst (1884:39) added: “With regard to polygama, 
Guen., a glance at his figure Noct. 3, pl. 16, f. 2, will 
convince any one that this species [blandula] could not 
have been intended. The primaries and secondaries are 
entirely different. The description accords with grynea, 
and the figure fits it as well as any species known to me.” 

Smith (1893) followed Hulst in placing both 
polygama sensu Guenée and alabamae as synonyms 
of grynea, and polygama sensu Grote as a synonym of 
blandula. Dyar (1903) apparently skirted the issue of 
Grote’s misidentification, and listed polygama as a full 
species with blandula as its synonym, and treated al- 
abamae as a full species. Hampson (1913) placed 
polygama as a synonym of grynea, both blandula and 
mira as synonyms of crataegi, and treated alabamae as 
a full species. In their monograph of the Nearctic 
Catocala, Barnes & McDunnough (1918b:40) treated 


polygama as a synonym of grynea, and both blandula 
and alabamae as full species, indicating: “It should be 
borne in mind that the ‘polygama Guenée, referred 
to by Lintner, Saunders, and others of the older au- 
thors is not the true species but probably what we 
have designated as blandula Hulst . .. Guenée’s figure 
of aolngenne is very poor but we do not see to what 
other species [i-e., grynea] it can be referred; it is cer- 
tainly not blandula.” McDunnough’s (1938) checklist 
followed Barnes and McDunnough’s taxonomy, and 
polygama has not been used as a species name since 
that time. Forbes (1954:336) placed polygama back 
under alabamae “(probably polygama Guenée)” as 
had Hulst, but Nearctic works after 1954 have listed 
polygama under the synonymy of grynea (e.g., Tietz 
1971, Hodges et al. 1983, Covell 1984, Poole 1989) as 
had Barnes & McDunnough. Note that some early 
20th century authors, notably R. Rowley, used 
polygama in error as the species name for mira (e.¢., 
Rowley & Berry 1910, cf. comments by Barnes & Mc- 
Dunnough 1918a:172-173; in his account of mira, 
Forbes 1954:335 indicated “polygama of American 
authors in large part, not Guenée”). 


Unquestionably, Grote’s initial misidentification of 


polygama, Hulst’s (1884) placement of both polygama 
and alabamae as synonyms of grynea, and the relative 
scarcity of specimen material for this group of closely 
related yellow-hindwinged species were all responsible 
for the volatile position of polygama in the 19th and 


early 20th century Nearctic literature. Some adults of 


this species group can be difficult to determine, but as 
a result of life history work during the first half of the 
20th century, and, especially, more recent studies by 
ourselves and colleagues (e.g., H. D. Baggett, W. A. 
Miller, D. F. Schweitzer, J. R. Slotten) it is now firmly 
established that alabamae, blandula, crataegi, grynea, 
mira, and pretiosa are all distinct species. The adults 
breed true, with multiple broods of each species hav- 
ing been reared ex ovo; many specimens of each 
species also have been reared ex wild larvae and the 
larvae of most are teens 

Guenée’s (1852) Fig. 2 on Plate 16 of polygama is 
clearly not blandula, and just as clearly is neither 
crataegi, mira, nor pretiosa. However, polygama i is also 
not gryned. Instead, Guenée’s figure of polygama is an 
acceptable albeit stylized render ing of alabamae, as 
suggested both by Forbes (1954) and originally by 
Hulst (1884:36) when he named blandula and cor- 


rected Grote’s misidentification. Guenée’s figure of 


polygama agrees with Grote’s description of and type 
of alabamae at the BMNH (accounting for sexual dif- 


terences: the alabamae type is a female, whereas the 
polygama tigure is a male), as well as to other speci- 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


mens of alabamae from along the Gulf Coast of the 
southern United States, especially Florida. Specific 
points of distinction include: (1) the forewing ground 
color of Guenée’s figure of polygama is warm light 
gray with brown shading, like alabamae, rather than 
deep cool greenish gray as in grynea; (2) the basal por- 
tion of ne forewing postmedial line and anal dash of 
polygama and alabamae are not profusely marked with 
rich, chocolate brown as in grynea; (3) the forewing 
antemedial and basal lines of polygama are present 
and distinct, as in alabamae, rather than wanting as in 
grynea; (4) the hindwing medial band of polygama and 
alabamae is thinner than in grynea, and not heavily 
suffused with dark scaling along the inner margin as in 
grynea; (5) the black hindwing margin is not as wide in 
polygama and alabamae as in grynea; (6) and the black 
hindwing margin is sharply discontinuous (separating 
sno a Cline peal dot) in polygama, as in alabamae, 

rather than continuous as is typical in grynea. 

The recently described Catocala charlottae Brou 
(1988) is the remaining taxon to which the name 
polygama could potentially apply. Brou’s original de- 
scription (1988:116, Figs. 1-4) distinguished charlot- 
tae in detail from alabamae, which is the only other 
similar species occuring with charlottae at its 
Louisiana type locality: charlottae differs from 
polygama on points 1 and 4-6 above in the same man- 
ner as grynea does from polygama. Moreover, the dark 
brown around the forewing anal dash is even more ex- 
tensive still in charlottae dhera j in grynea, normally con- 
tinuing into the anal margin. Thus, polygama does not 
refer to charlottae. 

In summary, since McDunnough (1938), the name 
polygama Guenée (1852) has been erroneously treated 
in the Nearctic Catocala literature as a junior synonym 
of grynea (Cramer 1780), rather than as a senior syn- 
onym of alabamae Grote (1875b). Until the ICZN is- 
sues a ruling on Case 3210, we maintain existing usage, 
retaining pol ygama under the synonymy of grynea. 


Catocala uxor Guenée, 1852. Hist. Nat. Spec. Gen. 
Lepid. 7:92 


The original description states “Amerique Septen- 
trionale. Coll. Bdv. Un °.” We have not located 
Guenée’s type, but there is no indication in the early 
Nearctic Catocala literature of confusion as to the ap- 
plicability of this name, and so we choose not to take 
any formal action. The type locality remains 
“Amerique Septentrionale.” The name wxor is a syn- 
onym of Catocala ilia (Cramer), and is preoccupied in 
the genus by uxor Hiibner (1788), a synonym of the 
Palearctic species Catocala nymphagoga Esper 
(1787). 


VOLUME 56, NUMBER 4 


Catocala viduata Guenée, 1852. Hist. Nat. Spec. 
Gen. Lepid. 7:400 


Guenée offered the name viduata in his Errata sec- 
tion (p. 400) as a nominal modification of J. E. Smith’s 
name vidua, a species which Guenée had misdiag- 
nosed earlier in his text (pp. 94-95). Hulst (1884) rec- 
tified this by proposing the replacement name maestosa 
Hulst for viduata (see discussion in Bames & McDun- 
nough 1918b:14 and Forbes 1954:325). The type local- 
ity of viduata is Georgia, [USA]. 


ACKNOWLEDGMENTS 


We are grateful to the following individuals for providing access 
to collections under their care, loans and/or photographs of type ma- 
terial, hospitality during our visits over the past decade, and related 
assistance: D. Azuma and J. Weintraub at the Academy of Natural 
Sciences (Philadelphia, Pennsylvania, USA); J. Miller, E. Quinter, 
and F. Rindge at the American Museum of Natural History (New 
York, New York, USA); W. Gall at the Buffalo Museum of Science 
(Buffalo, New York, USA); N. Penny at the California Academy of 
Sciences (San Francisco, California, USA); J. D. Lafontaine at the 
Canadian National Collection (Ottawa, Ontario, Canada); J. Rawlins 
at the Camegie Museum of Natural History (Pittsburgh, Pennsylva- 
nia, USA); J. Franclemont and J. Liebherr at the Comell University 
Insect Collection (Ithaca, New York, USA); W. Naessig at the 
Forschungsinstitut und Naturmuseum Senckenberg (Frankfurt, 
Germany); P. Parillo at the Field Museum of Natural History 
(Chicago, Illinois, USA); C. Brandao and A. Vasquez at the Museu 
de Zoologia (Sao Paulo, Brazil); W. Mey and W. Speidel at the Mu- 
seum fur Naturkunde of Humboldt University (Berlin, Germany); 
D. Bowers and S. Shaw at the Museum of Comparative Zoology of 
Harvard University (Cambridge, Massachusetts, USA); D. Fergu- 
son, D. Furth, M. Pogue, and R. Poole at the National Museum of 
Natural History (Washington D.C., USA); M. Honey and G. Martin 
at the Natural History Museum (London, United Kingdom); T. 
McCabe at the New York State Museum (Albany, New York, USA); 
G. McGavin and C. O'Toole at the Oxford University Museum of 
Natural History (Oxford, United Kingdom); H. Stein and H. Sturm 
at the Roemer- und Pelizaeus Museum and Hildesheim University 
(Hildesheim, Germany); C. Hauser at the Staatliches Museum fur 
Naturkunde (Stuttgart, Germany); M. Nuss at the Staatliches Mu- 
seum fur Tierkunde (Dresden, Germany); and P. Becker and H. 
Riemann at the Ubersee Museum (Bremen, Germany). 

For help with bibliographic matters, we also thank: C. Spawn at 
the Academy of Natural Sciences: M. Rose at the Atlanta History 
Center (Atlanta, Georgia, USA); R. Dirig at the Bailey Hortorium of 
Cornell University; B. Williams at the Field Museum of Natural His- 
tory; M. Epstein at the National Museum of Natural History; M. 
May at Rutgers University (New Brunswick, New Jersey, USA); and 
J. Harvey at the Natural History Museum. Many helpful comments 
on the manuscript were provided by R. Hodges (Eugene, Oregon, 
USA), D. Lafontaine, E. Quinter, D. Schweitzer (Port Norris, New 
Jersey, USA) and an anonymous reviewer. 


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Newman, London. pp. 983-1286. 


Received for publication 4 December 2001; revised and accepted 24 
July 2002. 


Journal of the Lepidopterists’ Society 
56(4), 2002, 265-271 


EARLY STAGES OF THE ENTOMOPHAGOUS METALMARK BUTTERFLY ALESA AMESIS 
(RIODINIDAE: EURYBIINI) 


P. J. DEVRIES AND C. M. PENz! 
Milwaukee Public Museum, 800 West Wells Street, Milwaukee, Wisconsin 53233, USA 


ABSTRACT. The immature stages of Alesa amesis are described in detail for the first time, and then compared to those of its sister genus, 


Eurybia. 


Additional key words: Eurybia, morphology, myrmecophily, caterpillar calls. 


The riodinid butterfly Alesa amesis (Cramer, 
1777) is a widespread and often locally common 
member of the tribe Eurybiini with a geographic 
range that includes Brazil, the Guyanas, Venezuela 
Colombia, Ecuador, and Peru. Recently we showed 
that A. amesis has an obligate association with Cam- 
ponotus femoratus (Fabricius, 1804) ants, and that 
the entomophagous caterpillars possess morphologi- 
cal and behavioral adaptations for feeding on Ho- 
moptera prey. These biological aspects are summa- 
rized briefly as follows. At one site in Amazonian 
Ecuador we found that female A. amesis oviposited 
only in the presence of C. femoratus ants tending ag- 
gregations of several genera of Membracidae or Ate- 
lionidae (Homoptera) that fed on six families of 
plants. Oviposition by A. amesis occurred either ad- 
jacent to aggregations of Homoptera, or directly on 
an individual nymph. Available evidence suggests 
that A. amesis caterpillars feed entirely on Ho- 
moptera nymphs, and that compared to other her- 
bivorous relatives, there has been an evolution of 
leg-length to accommodate their entomophagous 
diet. Greater comparative, behavioral, morphomet- 
ric and analytical details are provided in DeVries and 
Penz (2000). 

Given that there is little detailed information on 
early stages of most species of Eurybiini, and on Alesa 
in particular, here we present a detailed description of 
Alesa amesis early stages and compare them to species 
of their sister genus, Eurybia (Harvey 1987, Hanner 
1998). 


MATERIALS AND METHODS 


Field work was conducted at the La Selva Lodge, 
Garza Cocha, Sucumbios Province, eastern Ecuador in 
the upper Amazon Basin, 75 km E.S.E. of Coca 
(0°2950.3”S; 76°22’28.9”W). A detailed site descrip- 
tion is provided in DeVries and Walla (2001). 

Early stage material of A. amesis was field-col- 

' Adjunct professor at Programa de Pés-Graduagao em Biocién- 


cias, Pontificia Universidade Catélica do Rio Grande do Sul, Avenida 
Ipiranga 6681, Porto Alegre, Rio Grande do Sul 90619-900 Brazil. 


lected (caterpillars were first placed in Quinter’s so- 
lution, see protocol in DeVries 1997), then stored in 
70% alcohol, and later examined using light mi- 
croscopy. Except for the second instar, we examined 
all A. amesis early stages. Descriptions of caterpillar 
morphology follow the terminology of Peterson 
(1962), Cottrell (1984) and Stehr (1987). Preserved 
material of four Eurybia species was compared to 
first and fifth instar caterpillars, and pupae of A. ame- 
sis. This material included: fifth instar caterpillars of 
A. amesis, Eurybia patrona Weymer, 1874, E. elvina 
Stichel, 1910, E. nr. nicaeus (Fabricius, 1775) and E. 
lycisca (Westwood, 1851), and pupae of E. lycisca. 
Comparative differences and the sources of informa- 
tion are presented in Table 1, and Figs. 5-17. Com- 
parative voucher material of Alesa and Eurybia are in 
the Museo Nacional de Ecuador and the collection of 
DeVries. 


RESULTS 


Egg. (Fig. 1) (n = 2) Measurements: 0.83 mm wide, 
0.4 mm tall. White upon being laid, turning pale green 
within 24 hours; base broad, tapering gently towards 
apex; chorion heavily adorned with rounded sculptur- 
ing that is interconnected with numerous small tuber- 
cles. Egg bears little resemblance to lozenge-shaped 
eggs of the sister genus Eurybia, but it is reminiscent 
of the more distantly related Synargis (see illustrations 
in DeVries 1997). 

First instar. (Fig. 4) (n = 2) Head pale yellow- 
brown with short white plumose setae on anterior por- 
tions of epicranium and frons. Body white with short, 
white, finely barbed setae arising from brown pinnac- 
ula; dorsal pores on T1 and Al—A8; lateral body wall 
extended, flange-like, contacting substrate and con- 
cealing ventral side of body. Prothoracic shield pale 
yellow-brown, somewhat produced anteriorly and par- 
tially covering head; anterior margin of prothroacic 
shield with white, barbed, forward-projecting setae. 
Segments T1—T3 with distinct, pale brown dorsolateral 
crescent-shaped marks; thoracic legs white and dis- 
tinctly elongate. Prolegs white. Anal plate pale brown, 


266 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fics. 1-3. Alesa amesis early stages in nature. 1, Egg deposited directly on the abdomen of a membracid nymph. The overall form differs 
dramatically from the egg of Eurybia (illustrated in DeVries 1997:5). 2, Fifth instar caterpillar using thoracic legs to grasp its membracid prey. 
Note shortening of body segments to expose thoracic legs. 3, Pupa. Head oriented to the left. 


narrower than remaining body segments. No evidence 
for tentacle nectary organs or call production found in 
this instar. 

Second instar. No specimens of this instar were 
found. 

Third instar. (n = 3) Similar to fifth instar, and 
possessing bulb-shaped cuticular spinules covering 
body. As is general myrmecophilous riodinids (sum- 
marized in DeVries 1997), all third and subsequent 
instar caterpillars possessed a pair of tentacle nectary 
organs (hereafter TNOs) on segment A-8 (Figs. 2, 5, 
6, 8). As in other riodinid species the TNOs of A. 
amesis produced secretions only when solicited by at- 
tending ants (DeVries & Penz 2000). The ability to 
produce calls was also functional in this and all subse- 
quent instars. 

Fourth instar. (n = 6) Except for being smaller, in- 
distinguishable from fifth instar. Premolt duration 
from fourth to fifth instar 36 to 48 h. 

Fifth instar. (Figs. 2, 5-11) (n = 9) Head: black, 


anterior portion of epicranium and frons with 
minute tubercules and long simple setae; short 
simple setae distributed along entire surface of head; 
labrum narrow, exposing base of mandibles. Body: 
uniformly green except for light brown prothoracic 
shield and openings of tentacle nectary organs on A8 
(some fifth instars turned brown a few days after 
molting, and lost all traces of green). Lateral body 
wall extended, flange-like and with plumose setae, 
contacting substrate from A2-A9 and concealing 
prolegs and ventral side of body (Fig. 2); ventrolat- 
eral segmental areas reduced in T1-T3 and A1, tho- 
racic legs visible in lateral view (Fig. 2, see also 
DeVries & Penz 2000). Cuticle: covered with short 
spines set on broad, sclerotized bases armored with 
4-8 short points (most commonly with 6 points). 
Prothoracic shield: produced anteriorly and cover- 
ing head; in dorsal view, anterior margin of protho- 
racic shield with a distinct medial excavation and 3 
pairs of long plumose setae. Thoracic legs: white, 


VOLUME 56, NUMBER 4 


TABLE 1. 


267 


Comparative morphology of the early stages of Alesa amesis and Eurybia. All comparisons were done directly from preserved ma- 


terial unless indicated otherwise. Supplemental sources and notes are as follows: ' Horvitz et al. (1987); * Malicky (1970) discusses thickened in- 
tegument in larval Lycaenidae; * mandible examined in detail only in Ewrybia lycisca; * DeVries (pers. obs); ? DeVries and Penz (2000). Letters 
in first column correspond to details in Figs. 4-17, except for “body integument” (c), and “plantae” (m). 


First Instar—Fig. 4 


Alesa amesis 


Eurybia elvina ' 


a. Prothoracic shield in dorsal view 


b. Primary setae 
c. Dorsal pores 
d. TNOs 


longer and wider than head 
slightly covering head 
numerous, short, thick and finely barbed 
present on prothorax and Al—A8 
absent 


Fifth Instar—Figs. 5-17 


a. Body shape in lateral view 
b. Ventrolateral areas of body segments 


c. Body integument * 

d. Cuticular spines 

e. Frontal and adfrontal regions of head 
f, Distal segment of antenna 


g. Maxilla 
h. Mandible 


Alesa amesis 


distinctly humped at mid-length 

T1-T3 and Al conspicuously short, exposing 
legs; A2—AS8 elongated and hiding prolegs 

thicker than Eurybia and most other myrme- 
cophilous riodinids 

long, set on sclerotized base armored with 
4-8 points (most commonly 6) 

densely covered with thick, nub-like setae 


long 
comparatively small 
stout, with short teeth 


narrower and shorter than head 
not covering head 

sparse, long and thin 

absent 

present 


Eurybia patrona, E. elvina, E. lycisca, 
E. nicea 


not humped at mid-length 
largely uniform length across all body segments 


similar in thickness to most other myrme- 
cophilous riodinids 
short, without sclerotized base 


Eurybia patrona, E. niceaus, E. lycisca lack 
these setae; E. elvina with some scattered, 
slightly thicker setae 

short 

comparatively large 


i. Stemmata 
j- Prothorax in dorsal view 
k. Thoracic legs ° 


comparatively large 
covering head 


caterpillars 
|. Position of spiracles on Al 


m. Plantae comparatively broad 

n. Lateroseries of crochets crochets short and widely spaced 

o. TNOs externally stalked and armored 

p. Anal plate small and approximately rectangular 


posterior margin of A8 projected to 


enclose anal plate 
Pupa—Fig. 3 


a. Body shape stout 
b. Proboscis short 
c. Pupation site 


slightly darker at tarsi. Plantae of prolegs broad. 
Anal plate: rectangular, small, not well differenti- 
ated from rest of abdomen, and enclosed by postero- 
lateral expansions of AS. 

Pupa. (Fig. 3) (n = 7) Light to dark brown, elon- 
gate, tallest and widest anteriorly and tapering gradu- 
ally from head to cremaster. In ventral view, antennae 
terminate at posterior margin of A6, and proboscis ter- 
minates distally at posterior margin of T3 (for compar- 
ison, see Horvitz et al. 1987:517 fig. 4B, DeVries 
1997:137 fig. 40A). In lateral view, head and anterior 
portion of thorax resemble a miniature monkey face. 
T1 yellowish brown with an excavated anterior margin 


allometrically longer than other riodinid 


Alesa amesis 


uncovered pupae attached to plant stems 


slender, somewhat paddle-shaped, with long 
teeth ° 

comparatively small 

not covering head 

not differing allometrically from other riodinid 
caterpillars 


centered and slightly above the spiracular line near anterior margin and below the spiracular 


line 
comparatively narrow 
crochets long and densely packed 
externally an un-stalked slit 
large and oval, posterior margin of A8 not 
projected 


Eurybia elvina ', E. lycisca 


slender 

elongate, extending beyond cremaster 

typically concealed within sheathing stems of 
host plant }4 


and covering head in dorsal view; thoracic spiracle red. 
T2-T3 with large white to green crescent mark. Pupa 
with skirt on A4Q—A10 that projects ventrally and flares 
over substrate; ventral side concave from T3 (approxi- 
mately) to Al0 to adjust for shape of pupation sub- 
strate. Silk girdle across Al; Al and A8 conspicuously 
shorter than A2—A7; AY reduced. Cremaster broad, 
longer than other abdominal segments, and slightly 
curved ventrally. Duration of pupa 15-17 days (n = 3; 
2 males, 1 female). Pupation in nature occurred on 
small stems at base of plants associated with 
Camponotus femoratus colonies, and pupae were at- 
tended by ants. 


To eS 


a AS SOP 


Fic. 4. First instar of Alesa amesis. Note size of prothoracic 
shield (a), primary setae (b), and dorsal pores (e). 


DISCUSSION 


Previously the only available detailed description of 
Eurybiini early stage morphology was that of Eurt rybia 
elvina by Horvitz et al. (1987). The present study 
shows that caterpillars and pupae of A. amesis may 
differ considerably from Eurybia (Table 1, Figs. 
4-17). For ex ample, although all instars of Ewrybia 
elvina apparently possess TNOs (Horvitz et al. 1987) 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


these organs are absent in first instar A. amesis. Other 
differences among A. amesis and Eurybia caterpillars 
include body shape, cuticular spines, relative thoracic 
and abdominal leg size, crochets, stemmata, anten- 
nae, and maxilla (Figs. 5-17). Traits like elongated 
thoracic legs, broad proleg plantae and the ventrolat- 
eral shorter tine of segments T1—-3 and Al in A. amesis 
caterpillars may reflect their entomophagous habit 
since they potentially facilitate curling of the body 
during prey capture and feeding. Other characteris- 
tics of A. amesis caterpillars may be due to their form- 
ing symbioses with ants (e.g., long cuticular spines in 
sclerotized, armored bases; thickened body integu- 
ment; armored, stalked TNOs). However, the evolu- 
tionary basis and adaptive nature of such traits remain 
uncertain. 

Elsewhere we have described the substrate-borne 
calls of A. amesis caterpillars, noting that the mecha- 
nism for call production within the Eurybiini was un- 
known (DeVries & Penz 2000). Recently Travassos et 
al. (in press) presented evidence suggesting that 
Eurybia elvina produce a substrate-bome call by grat- 
ing cervical membrane “teeth” against hemispherical 
protuberances on the surface of the head. Our exami- 
nation of A. amesis caterpillars with optical mi- 
croscopy revealed a cervical membrane similar to that 
described by Travassos et al. (in press) for E. elvina— 
the membrane is armored with “teeth” and bears 
small setae and rounded protuberances. Moreover, 
we note the presence of rounded sclerotized areas in 
the dorsolateral portion of the cervical membrane in 
A. amesis and E. lycisca, E. patrona, E. lycisca, E. nv. 
nicaeus and E. elvina (indicated by arrows in Fig. 10). 
The function of these rounded sclerotized areas is un- 
clear, but their potential role in caterpillar call pro- 
duction warrants further investigation because they 
occur in proximity to where epicnaniel granulations 
are well developed. 

Although brief in scope, we hope this study will 
stimulate comparative life history work on other 
species of Eurybiini to further our understanding of 
the biology and evolution of this unusual group of rio- 


dinids. 


ACKNOWLEDGMENTS 


We thank the La Selva Lodge, its staff, and particularly Eric 
Schwartz for providing strong field support for this and other stud- 
ies. Comments and suggestions by L. Gall, R. Hill, E. Youngsteadt, 
D. Wagner and A. Warren improved the manuscript. We thank 
Museo de Ciencias Naturales (Quito, Ecuador) for assistance in this 
and other studies. We gratefully acknowledge the help H. Greeney, 
R. Hill and N. Gerrardo for help in caring for early stages. This 
study was supported in part by NSF DEB 00-96241. We dedicated 
this paper to the late Tommy Flanagan, Joe Henderson, and Vinicius 
de Morais. 


269 


VOLUME 56, NUMBER 4 


aly. 
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Details of Alesa amesis fifth instar caterpillar. Figs. 5-10 represent the same individual; Fig. 11 dissected from a cast skin. Let- 


Fics. 5-11. 
ters correspond to traits listed in Table 1. 5, Dorsal view of caterpillar, head oriented to the right. 6, Tentacle nectary organ. 7, Crochets of third 
abdominal proleg. 8, Caterpillar in ventrolateral view. 9, Detail of the caterpillar thorax in ventrolateral view, insets show body and head spines. 


10, Detail of caterpillar head and cervical membrane (head setae and body spines omitted), arrows indicate the round sclerotizations present 
on the cervical membrane. 11, Outer (top) and inner (bottom) views of left mandible. Letters correspond to details in Table 1, except for “body 


integument’, and “plantae”. 


bo 
>I 
> 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 12-17. Details of Eurybia patrona and E. lycisca fifth instar caterpillars. Figs. 12-16 represent the same individual E. patrona (Barro 
Colorado Island, Panama); Fig. 17, E. lycisca mandible drawn from a cast skin (Parque Nacional Corcovado, Costa Rica). 12, Dorsal view of 
caterpillar, head oriented to the right. 13, Tentacle nectary organ. 14, Crochets of third abdominal proleg. 15, Caterpillar in ventrolateral view. 
16, Detail of the caterpillar thorax in ventrolateral view, inset shows a body spine. 17, Outer view of E. lycisca left mandible. Letters correspond 


to details in Table 1, except for “body integument’, and “plantae”. 


LITERATURE CITED 


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DeVries, P. J. 1997. The butterflies of Costa Rica and their nat- 
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ton. 

DeVries, P. J. & C. M. Penz. 2000. Entomophagy, behavior, and 
elongated thoracic legs in the myrmecophilous Neotropical but- 
terfly Alesa amesis (Riodinidae). Biotropica 32:712-721. 


DeVries, P. J. & T. R. WALLA. 2001. Species diversity and commu- 
nity structure in neotropical fruit-feeding butterflies. Biol. J. 
Linn. Soc. 74:1—-15. 

HANNER, R. H. 1998. Taxonomic problems with phylogenetic solu- 
tions derived from the integration of biochemical, morphologi- 
cal and molecular data. Unpublished Ph.D. Dissertation, Uni- 
versity of Oregon, Eugene, Oregon. 

Harvey, D. J. 1987. The higher classification of the Riodinidae (Lep- 
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Austin, Texas. 


VOLUME 56, NUMBER 4 


Horvitz, C. C., C. TURNBULL & D. J. Harvey. 1987. Biology of im- 
mature Eurybia elvina (Lepidoptera: Riodinidae), a myrme- 
cophilous metalmark butterfly. Ann. Entomol. Soc. Am. 
80:513-519. 

PETERSON, A. 1962. Larvae of insects. part I: Lepidoptera and 
plant infesting Hymenoptera. Edwards Bros, Columbus. 

Maticky, H. 1970. New aspects of the association between ly- 
caenid larvae (Lycaenidae) and ants (Formicidae, Hy- 
menoptera). J. Lep. Soc. 24:190-202. 


TAL 


STEHR, F. W. 1987. Order Lepidoptera, pp 288-305. In Stehr, F. 
W. (ed.), Immature insects. Kendall/Hunt, Iowa. 

Travassos, M., P. J. DEVRIES & N. E. PreRCcE. In press. A novel or- 
gan and mechanism for larval sound production in butterfly 
caterpillars: Ewrybia elvina (Lepidoptera: Riodinidae). Trop. 
Lepid. 


Received for publication 4 March 2002; revised and accepted 30 
May 2002. 


Journal of the Lepidopterists’ Society 
56(4), 2002, 272-276 


THE EFFECTS OF SEASON, HOST PLANT PROTECTION, AND ANT PREDATORS ON THE 
SURVIVAL OF EUMAEUS ATALA (LYCAENIDAE) IN RE-ESTABLISHMENTS 


EILEEN M. SMITH! 
Department of Environmental Studies, F lorida International University, Miami, Florida 33199, USA 


ABSTRACT. The primary purpose of this study on Eumaeus atala, the atala butterfly, was to determine which factors influence larval sur- 
vival during re-establishment of atala butterfly populations. An inexpensive protective cover of fabric netting over the host plants at the re- 
establishment site was found to have a positive effect on the number of larvae that survived to pupation. Season was also found to have an effect 
on the number of re-established larvae that survived to pupation. Significantly more larvae survived to pupation during the wetter summer sea- 
son than during the drier winter season. This suggests that future attempts to re-establish the atala should take place in the summer and should 
consider the use of protective netting over host plants. In the course of this study, the mortality of atala eggs was found to be high, and two new 


ant predators of atala eggs were found. 


Additional key words: coontie, cycasin, insect. 


The atala, Eumaeus atala Poey, is a hairstreak but- 
terfly (Lycaenidae, subfamily Theclinae) with a tumul- 
tuous history. It was once considered to be extinct 
throughout south Florida due to the exploitation of its 
sole native larval host plant, but the atala has since 
made a comeback. There are close to 50 lycaenid 
species in the West Indies and south Florida (Smith et 
al. 1994). Female atala lay whitish-yellow eggs in clus- 
ters of up to thirty eggs which are usually deposited on 
fresh leaves of cycad species or on their reproductive 
cones. Atala larvae are a rusty-red color, with seven 
pairs of canary yellow spots on the dorsal side and 
short black setae. Their bright coloring is aposematic, 
as larvae and adults contain cycasin (Bowers & Larin 
1989, Nash et al. 1992). Spiders, for example, who find 
the atala in their webs, avoid them (Hubbuch 1991). 
The atala apparently concentrate the secondary com- 
pounds of their larval host plant, the Florida coontie, 
Zamia pumila (Zamiaceae), in their bodies. 

Eumaeus atala had a historical range of Dade, Mon- 
roe, and Broward counties in Florida. It also ranged 
throughout Cuba, and into the Bahamas (Clench 1943). 
In Florida, the atala’s current range includes Dade, 
Broward, Monroe, Palm Beach, Indian River and St. 
Lucie counties (Culbert 1995). It also still occurs in 
Cuba, the Bahamas, and Cayman Brac (Hammer 1995). 

Early scientific references to the atala describe the 
species as abundant in south Florida and Cuba (Scud- 
der 1875, Schwartz 1888, Healy 1910, Grossbeck 
1917). By the mid-twentieth century, the atala was 
considered rare or extinct in Florida by lepidopterists 
(Ford 1946, Klots 1951, Young 1956, Funk 1966, Raw- 
son 1961). In 1979, on Key Biscayne, Florida, Miami- 
Dade county naturalist Roger Hammer found a thriv- 
ing colony of Eumaeus atala (Roger Hammer pers. 
com.). Hammer attempted re-establishments, many of 
which were successful. 

' Current affiliation: Department of Comparative Sociology, 


Florida International University, University Park Campus, Miami, 
Florida 33199. E-mail: esmithO1@fiu.edu 


Atala multiplied at various locations in South 
Florida from Coral Gables to Florida City (Landolt 
1984). “The atala has made a spectacular recovery and 
is now found in urban and natural areas around Fort 
Lauderdale and Miami” (Emmel & Minno 1993). The 
Key Biscayne colony, however, vanished for unknown 
reasons around 1991. 

The main goal of this project was to conduct an atala 
re-establishment and to examine three questions. The 
first was to find out whether season would have an ef- 
fect on larval and pupal mortality in a re-establishment. 
The second question was whether protecting atala host 
plants would increase survivability of larvae. It was hy- 
pothesized that if netting were placed over the host 
plants, more of the translocated larvae would survive 
compared to larvae on unprotected plants. The third 
question was are there predators of atala larvae, eggs 
and pupae, despite their cycasin content. This is im- 
portant information because despite the fact that re- 
establishments appear to have preserved the atala in 
Florida, few details are known about what factors en- 
hance success in re-establishment. 

All of the study sites for this project were in Bear 
Cut Preserve, Crandon Park on Key Biscayne, Florida. 
Restoration of habitat was necessary prior to attempt- 
ing atala re-establishment. The coastal maritime ham- 
mock where the study took place had been damaged 
by fire and exotic plant species (Doren et al. 1991). 
Few coontie plants remained in Crandon Park. Thirty- 
one coontie plants were purchased and planted in the 
re-establishment areas to improve the atala habitat 
prior to this project. 


MATERIALS AND METHODS 


Atala translocation to Key Biscayne began in Jan- 
uary of 1999. Miami's Fairchild Tropical Garden 
donated all larvae and pupae used in the re- 
establishment. Fairchild was chosen as a source since 
it has the most consistently strong atala population in 
the county and because its colony originated from the 


VOLUME 56, NUMBER 4 


1980’s human-established atala colonies. The number 
of larvae used in releases for this project and the tim- 
ing of releases themselves were always dependent on 
the population size of the Fairchild colony. 

Atala larvae were all close to the same age at translo- 
cation, determined by measurement to be in their sec- 
ond instar. They were removed from the colony at 
Fairchild Tropical Gardens and placed on Z. pumila at 
four different sites on Key Biscayne with similar plant 
cover, topography, accessibility to host plants, and light 
levels. In addition to the larvae placed at these sites in 
Crandon, corresponding larvae were placed at an off- 
site location. Larvae were placed in three different 
treatment situations. 

The three different treatments to the larvae were as 
follows. In Treatment #1, atala larvae were released 
and placed on Z. pumila in Crandon Park in a “nat- 
ural” release with no treatment. Larvae had no protec- 
tion from predators, except for their natural defenses. 
In Treatment #2, larvae were placed on Z. pumila in 
Crandon Park, then covered with a fine green nylon 
netting fabric that was tightly tied at the base of the 
plant. Holes in the mesh were 10 mm. Upon pupation 
of the atala, this netting was removed. In Treatment 
#3, the larvae were reared in captivity off-site on fresh- 
cut Z. pumila fronds under a high level of protection 
on a screened porch without temperature control. This 
third treatment gives an indication of how survival of 
atala in captive rearing compares to survival in the 
field. 

The attempted re-establishments in Crandon Park 
and the off-site captive rearing experiments were re- 
peated three times throughout the year: Winter/Dry 
season, Transitional/Spring season and Summer/Rainy 
season (see Table 1). 

Length of residency was used as a measure of sur- 
vival for larvae. If a larva was no longer on the host 
plant or was not discovered pupating on other plants 
within 1 m of the host plants, it was considered to have 
died, even if the actual “corpse” was not discovered. 
Although larvae sometimes leave the host plant to pu- 
pate, they tend to do so in clusters and are easily found 
usually within 1 m of the host plant regardless of 
whether netting is present. This was reinforced in my 
experiment at an off-site location on a screened porch 
where, despite the lack of netting around coontie, 76% 
of atala larvae did not venture farther than 0.5 m from 
their point of translocation to pupate. The other 24% 
of larvae crawled up to 1 m away, but never to a 
greater distance when sufficient coontie was available, 
despite there being no barriers to prevent them from 
doing so. At both the park and the offsite location, 
while there were other plants surrounding the translo- 


273 


TABLE 1. Experiments #1-3: an explanation of atala re- 
establishments in the dry/winter season, transitional season, and 
summer/wet Season. Experiment #1 is the Dry/Winter season. Ex- 
periment #2 is the Transitional Season. Experiment #3 is the 
Wet/Summer Season. 


Total # Sites 
Date Experiment oflarvae Treatment released 
1/15/99 #1 59 none ILD. 8), 4 ts) 
1/28/99 #1 59 captivity ex situ 
2/16/99 #1 59 netting IL, 2, Bh, 4b fs) 
4/9/99 #2 21 none ® 
5/5/99 #2 BD captivity ex situ 
5/5/99 #2 21 netting 2 
6/30/99 #3, 54 none Les 
6/30/99 #3 54 captivity ex situ 
6/30/99 #3 54 netting ).5) 


cation sites, the only coontie available were those that 
were part of the experiments. 

Pupal mortality was distinguished by presence and 
condition of the pupae. When an atala butterfly 
emerges naturally, there is a cleanly consistent ecdys- 
tal slit in the cuticle. If the pupa has been opened by 
outside force, this is easy to discern. If the pupa van- 
ished, it was assumed to have been taken by a predator 
unless discovered in its entirety on the ground beneath 
the plant on which it pupated. 

Ideally, Treatment #1, Treatment #2, and Treatment 
#3 would all have been initiated simultaneously at the 
beginning of each of the three seasons. However, lar- 
vae were not always available in sufficient numbers at 
Fairchild, so some of the differing larval treatments in 
the same experiment were separated by 2-4 week time 
spans (still within the same season). 

The four sites in Crandon Park, sites #1, 2, 3, and 5 
were assumed to be similar to one another in plant 
cover, light level and topography and were used as 
replicates (Site 4 was removed from the study prior to 
its onset). Sites 6-13 were off-site captive-rearing sites 
assumed to be similar to one another and used as 
replicates. 

To address the hypothesis that netting improved the 
ability of larvae to survive to pupate and the question 
of whether season impacted larval survival, analyses of 
variance were performed by treatment and season. Pu- 
pal survival to emergence was also examined in these 
ANOVAs. Tukey’s post hoc test was also performed. 


RESULTS 


There was no interaction between treatment and 
season on larval survival to pupation, F (4, 15) = 2.30, 
p = 0.107. There was also no interaction between 
treatment and season on survival to emergence, F (4, 
15) = 0.35, p = 0.842. 


bo 
~l 
aN 


TABLE2. Main effects of season and treatment on atala pupation 
and emergence. 


Analyses of variance 


Main effects of df F Significance 
TREATMENT on % pupating — (2, 15) 16.13 p < 0.001 
TREATMENT on % emerging (2, 15) 3.08 p< 0.076 
SEASON on % pupating (2, 15) 6.86 p < 0.008 


There was a main effect of treatment on pupation, F 
(2, 15) = 16.13, p < 0.001 (Table 2). Tukey's post hoc 
test at a 5% significance level demonstrated that the 
mean percentage of atala pupating with protective net- 
ting (62.7%) was significantly greater than the mean 
percentage pupating without netting (17.1%) (Table 3). 

There was also a marginally significant main effect 
of treatment on emergence, F (2, 15) = 3.08, p < 0.076 
(Table 2). Tukey’s post hoc test at a 5% significance 
level revealed that the mean percentage of atala that 
emerged from their pupae successfully on plants 
where netting had been applied earlier on (96.9%) was 
significantly higher than the mean percentage that 
emerged from their pupae without netting ever having 
been used (60.8%) (Table 3). 

As predicted, there was a significant main effect of 
season, F (2, 15) = 6.86, p < 0.008 (Table 2). Tukey's 
post hoe test indicated that Season #3/Summer had a 
significantly higher percentage (56.8%) of pupation 
than Season #1/Winter (33.2%) (Table 3). 

Few predators of the atala in any of its life stages have 
ever been reported. In the course of this study, several 
ant species were found to be major predators on atala in 
the egg stage, and one species was found to enter pu- 
pae. The first ant species noted as an egg predator was 
Camponotus abdominalis var. floridana. This is a com- 
mon native ant in Florida that often infests dwellings. 
Also called “bull dog” ants, they are known to feed on 
honeydew and insects (Smith 1972). In prior lab feeding 
trials, atala adults and cycasin were considered deter- 
rents to C. abdominalis (Bowers & Larin 1989). 

The same species of ant was observed at Site #2 in 
Crandon on 4 June 1999 tearing apart atala eggs where 
19 eggs had been laid. At the same time, an atala but- 
terfly was fluttering around the plant in a pattern typi- 
cal of an egg-laying female. The butterfly got very 
close to the frond where the ant was eating eggs, and 
beat at the ant with its wings. The ant grabbed an an- 
tenna of the atala butterfly and pulled. There was a 
fierce struggle, and the butterfly fell to the ground. 
The atala beat its wings on the ground, momentarily 
unable to fly, then flew away quickly. This unusual be- 
havior appeared to be very purposeful on the part of 
the atala, as if it were attempting to drive the ant away 
trom its offspring. 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


TABLE 3. Tukey's post hoc test at 5% level: mean percentage of 
atala pupation and emergence by treatment group and by season. 


Mean Mean 
% pupating: % emerging: 
BY TREATMENT GROUP 
#1 Group (with No Treatment)* 17.1% 60.8% 
#2 Group (with Medium 62.7% 96.9% 
Treatment/Protection Level)” 

#3 Group (with High Protection)® 54.7% 96.7% 
Significant difference: 
Group #1* vs. #2? p < 0.001 p < 0.046 
Group #2, vs. #3: N.S. N.S. 
Group #1* vs. #3° p < 0.001 p < 0.047 
BY SEASON 
#1 Season! 33.2% 82.3% 
#2 Season® 55.6% 84.6% 
#3 Season! 56.8% 88.2% 
Significant difference: 
Season #14 vs. #3! p < 0.009 nS. 
Season #14 vs. #2° n.s. nS. 
Season #3! vs. #2° N.S. N.S. 


“In situ without netting on plants 
'In situ with netting on plants 

“Ex situ in captivity (outside cages) 
4Winter/Dry Season 

© Transitional/Spring Season 

: Summer/Rainy Season 

n.s. = no significance, p > 0.05. 


It took a Camponotus ant very little time to find 
atala eggs. On 30 August 1999, a single ant was ob- 
served at Site #2 at 13:32 on a coontie frond while an 
atala was laying eggs. The ant came near enough to the 
six eggs to attack them, but did not, then tumed 
around and left the plant. Ten minutes later, another 
ant (possibly the same one) appeared and began to 
tear a hole in the eggs. Within eighteen minutes of the 
egg-laying event, three other ants of the same species 
arrived and began a new attack on the eggs, until all six 
were destroyed. 

On 5 July 1999 a large ant (Pseudomyrmex mexi- 
canus) was observed attacking atala eggs at Site #2 
where 25 eggs had been laid. Nine undisturbed pupae 
were also present on the same plant. Pseudomyrmex 
mexicanus is native and found throughout the eastern 
United States (Smith 1972). 

Another egg predator was observed during a survey 
at Rockdale Pineland in Miami on 3 October 1999. 
Wasmannia auropunctata (Roger) spent 15 min punc- 
turing a small hole in an egg before the ant was col- 
lected. Commonly known as the “little fire ant,” W. au- 
ropunctata is a neotropical ant introduced into Florida 
(Smith 1972). 

For several weeks, pupae would occasionally be 
found in Crandon Park with a perfectly round 0.1 em 
diameter hole in the outer layer. Several times, upon 
examination, tiny ants (Monomorium floricola) spilled 


VOLUME 56, NUMBER 4 


Fic. 1. A female Ewmaeus atala (atala butterfly) depositing eggs on a Zamia pumila (coontie) frond. Photograph by Robert Schroeder (used 


with permission). 


out of the holes. Monomorium floricola is an intro- 
duced ant from Africa or Asia, known to feed on in- 
sects (Smith 1972). These ants may have been oppor- 
tunists who went into holes made by a parasite. 

In Bear Cut Preserve, Crandon Park, ants appeared 
to be a major cause of egg mortality and had a serious 
effect on atala survival. Ants left visual evidence be- 
hind in the form of characteristic broken and torn 
apart eggshells. Ants were also observed, less com- 
monly, carrying eggs away from the plant. Over 700 
atala eggs were observed to have been laid in Crandon 
Park. Of these, 131 were destroyed in a manner that 
implicated ants and at least the same number of eggs 
simply vanished. 


DISCUSSION 


The results of this re-establishment provide input 
for a plan of successful re-establishment of the atala. 
The survival of translocated larvae was greatly en- 
hanced by a simple and inexpensive protective netting 
treatment that affords extra protection in the vulnera- 
ble larval stage. Survival of these protected in situ lar- 
vae was equivalent to the survival of highly protected 
captive-reared larvae. It is therefore recommended 
that atala re-establishments use protective netting to 


cover larvae and plants until at least pupation. Re- 
establishment utilizing pupae rather than larvae 
should also be tested, since once atala pupate, survival 
is quite high, even without netting protection. 

As soon as all larvae in the experiments had pu- 
pated, the netting was removed, so the use of netting 
might not necessarily be expected to affect the per- 
centage of atala that survived from pupation to emer- 
gence. Despite this, there was marginal significance, 
with more butterflies successfully emerging from pu- 
pae that had been protected by netting in their larval 
stage. Further testing should be conducted to deter- 
mine how the netting utilized in the larval stage may 
enhance future pupal survival. 

Larval survival was highest in the summer. The 
summer season is also the wettest. It is a logical out- 
come since during the rainy season, there are more 
fresh coontie fronds available, which are the favored 
food of early instars. During the dry season, the 
young atala caterpillars must work harder to survive, 
scraping at the underside of old, tougher fronds. Re- 
establishments of the atala appear to have a much 
greater chance of success when conducted in the sum- 
mer rainy season. 

Egg protection must be factored into the re- 


bo 
=I 
o>) 


establishment equation. The impact of native and ex- 
otic ant species on the atala requires a detailed exami- 
nation. Finer netting, placed over eggs as soon as pos- 
sible, may discourage these and other predators. 
Re-establishment of the atala into habitat where it 
was once common, but has been locally extirpated, is 
possible with a limited time commitment and limited 
financial investment, and the results appear to be very 
promising. The re-established atala colony on Key Bis- 
cayne, with no translocations having been done since 
June of 1999, was still abundant at the time of the last 
survey by this researcher in November of 2001. 


ACKNOWLEDGMENTS 


Thanks go to the Florida International University Tropical Biol- 
ogy Grant Program and the Marjory Stoneman Douglas Biscayne 
Nature Center ep financial help in this endeavor. Dr. Suzanne Kop- 
tur generously shared her expertise as major advisor and in ant iden- 
tification. Dr. Alice Clarke and Dr. Thomas Pliske were very helpful 
committee members. Thanks also go to Fairchild Tropical Gardens 
and Director of Collections, Chuck Hubbuch for assistance in 
procuring atala larvae. Dr. Robert Schroeder was extremely gener- 
ous in lending his brilliant photographs of the atala. The staff of 
Crandon Park in Miami-Dade County and Miami-Dade Natural Ar- 
eas Management facilitated the field work. Finally, this project could 
not have occurred without interviews from Roger Hammer and sup- 
port from Professor Douglas D. Molash. 


LITERATURE CITED 


Bowers, M. D. & Z. Larin. 1989. Acquired chemical defense in 
the lycaenid butterfly Eumaeus atala. J. Chem. Ecol. 
15:1133-1147. 

CLENCH, H. kK. 1943. The Lycaenidae of the Bahama Islands. 
PSYSA 49:52-60, 

CULBERT, D. 1995. Florida coonties and atala butterflies. Pam- 
phlet. Indian River County Extension Service, Vero Beach, 
Florida. 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


DoreEN, B., G. GANN, D. HARDIN, A. HERDON, G. KNIGHT, J. 
MAGuIRE, E. MENGES, J. KING, J. STENBERG & J. SNYDER. 
1991. Crandon Park natural areas protection plan. Miami: 
Metro Dade County Park and Recreation Dept. paper in con- 
junction with the Nature Conservancy and Fairchild Tropical 
Garden. 

EMMEL, T. & M. C. MINNO. 1993. Butterflies of the Florida Keys. 
Scientific Publishers, Gainesville, Florida. 

ForD, E. B. 1946. Butterflies. Collins, London. 

FuNK, R. 1966. Record of Eumaeus atala from the florida keys. J. 
Lepid. Soc. 20:216. 

GrROSSBECK, J. A. 1917. Insects of Florida: Lepidoptera. Bull. Am. 
Mus. Nat. Hist. 82:1—2. 

Hammer, R. 1995. The coontie and the atala hairstreak. The Pal- 
metto 15:3-5. 

HEALy, J. L. 1910. The larvae of Eumaeus atala florida. Ent. News 
21:179-180. 

Hupsucn, C. 1991. Wings of a dilemma; the atala butterfly. 
Fairchild Trop. Gar. Bull. 46:20-23. 

Kuors, A. 1951. A field guide to the butterflies. Houghton Mifflin, 
Company, Boston. 

LANDOLT, P. J. 1984. The Florida atala butterfly, Eumaeus atala 
florida, Fla. Entomol. 67:570-571. 

Nasu, R. J., E. A. BELL & P. R. ACKERY. 1992. The protective role 
of cycasin in  cycad-feeding Lepidoptera. PYT'CA 
31:1955-1957. 

Rawson, G. 1961. The recent rediscovery of Eumaeus atala in 
south Florida. J. Lepid. Soc. 15:237-244. 

ScHWakZ, E. A. 1888. Notes on Eumaeus atala. Insect Life 
1:37—-40. 

SCUDDER, S. H. 1875. The structure and transformation of Eu- 
maeus atala. Boston Soc. of Nat. Hist. Memoirs 2:413-419. 
SmiTH, D., J. MILLER & L. MILLER. 1994. The butterflies of the 

West Indies and south Florida. Oxford University Press, Oxford. 

SmiTH, M. R. 1972. House infesting ants of the eastern United 
States. Technical Bulletin 1336. United States Department of 
Agriculture. 

YOUNG, F. 1956. Notes on collecting Lepidoptera in southern 
Florida. J. Lepid. Soc. 9:204-212. 


Received for publication 10 July 2001; revised and accepted 15 Feb- 
ruary 2002. 


Journal of the Lepidopterists’ Society 
56(4), 2002, 277-285 


A NEW SPECIES OF EPIBLEMA PREVIOUSLY CONFUSED WITH E. TRIPARTITANA (ZELLER) AND 
E. INFELIX HEINRICH (TORTRICIDAE) 


DONALD J. WRIGHT 
3349 Morrison Ave., Cincinnati, Ohio 45220-1430, USA 


ABSTRACT. Epiblema glenni, new species, is described from 36 adult specimens (29 d, 7 2). This species frequently has been misiden- 
tified as E. tripartitana (Zeller), its closest congener, from which it differs in details of forewing maculation. Records of E. glenni from seven 
states suggest a range extending from central Missouri to western North Carolina and north to central Michigan. A rather long history of con- 
fusion involving E. glenni, E. tripartitana, E. infelix Heinrich, and E. scudderiana (Clemens) is reviewed. Adults and genitalia (6 and 2) of all 


four taxa are illustrated. 


Additional key words: Olethreutinae, Eucosmini, Epiblema scudderiana. 


About one-third of the 40 recognized species of 
Epiblema (Hiibner) in North America feature a pre- 
dominantly dark forewing with a prominent whitish 
spot at the middle of the dorsal margin. One of the 
more conspicuously marked members of this group is 
E. tripartitana (Zeller), whose spot extends from the 
dorsal margin to the costa, forming a broad transverse 
band that completely separates the dark basal and ter- 
minal portions of the forewing. 

While surveying remnant prairie habitat in Adams 
County, Ohio, I encountered a new species of Epi- 
blema with forewing markings that are similar to yet 
clearly distinguishable from those of E. tripartitana. 
Investigation turned up additional representatives of 
this species, collected in the 1960's by Murray O. 
Glenn in Putnam County, Illinois, and mixed in two 
public collections with specimens of E. tripartitana. I 
could find no consistent differences in male genitalia 
between the new species and tripartitana, and prepa- 
rations of neither matched Heinrich’s (1923, Fig. 270) 
illustration of the latter species. The reason for this 
discrepancy proved to be a misinterpretation by Hein- 
rich of several specimens from Florida. The holotype 
of tripartitana is a female from Dallas, Texas. Hein- 
rich’s (1923:146) treatment of this species refers to a 
series of specimens from Texas and Florida, and he il- 
lustrated the genitalia of a male from Cocoanut Grove, 
Florida. Examination of that specimen and of a num- 
ber of similarly marked specimens from nearby 
Florida localities revealed that they are not conspecific 
with tripartitana. They agree in both male and female 
genitalia with Epiblema scudderiana (Clemens). Their 
forewing coloration is sufficiently near that of triparti- 
tana to cause the aforementioned confusion, but the 
basic forewing pattern does match that of scudderiana. 
Moreover, I found a reared specimen of this species 
from Savannah, Georgia, labeled “Goldenrod stem 
borer,” suggesting that it shares a common life history 
with scudderiana. For these reasons, I decided to treat 
the Florida specimens as representing a darkly marked 
population of scudderiana. Infraspecific variation in 
scudderiana is discussed below. 


The proper application of the name tripartitana was 
confirmed by examining the holotype. In studying ma- 
terial from various collections I discovered that the 
new species also has been confused with E. infelix 
Heinrich, so an account of infelix is included for diag- 
nostic purposes. 


MATERIALS AND METHODS 


I examined specimens from the following institu- 
tional and personal collections: American Museum of 
Natural History (AMNH), George J. Balogh (GJB), 
University of Connecticut (UConn), Loran D. Gibson 
(LDG), Illinois Natural History Survey (INHS), Uni- 
versity of Louisville (UL), Michigan State University 
(MSU), Mississippi Entomological Museum (MEM), 
Museum of Comparative Zoology (MCZ), Ohio Lepi- 
dopterists (OL), United States National Museum of 
Natural History (USNM), James R. Wiker, and Don- 
ald J. Wright (DJW). Other cited collectors are abbre- 
viated as follows: Julian P. Donahue (JPD), E. H. Metz- 
ler (EHM), Murray O. Glenn (MOG). Line drawings 
were made with the aid of a Ken-A-Vision micropro- 
jector (Model X1000-1). Terminology regarding 
forewing pattern elements follows Brown and Powell 
(1991). Forewing length was measured from base to 
apex, including fringe, and the number of specimens 
supporting a particular statistic is indicated by n. 


SYSTEMATICS 


Epiblema tripartitana (Zeller) 
(Figs. 2, 5, 12, 16) 


Paedisca tripartitana Zeller 1875:308, pl. 9, Fig. 39 
(forewing). 

Eucosma tripartitana Fermald [1903]:459; Barnes and 
McDunnough 1917:171. 

Epiblema tripartitana (not Zeller 1875) Heinrich 
1923:146; Kimball 1965:260. 

Epiblema tripartitana McDunnough 1939:48; Powell 
1983:35; Miller 1987:57 (wings, 6 & ° genitalia). 
Holotype. °: Dallas, Tex., Boll; MCZ Type No. 

14337. 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 1-6. 1. E. glenni, holotype male, Adams Co., Ohio. 2. E. tripartitana, male, Ithaca, New York. 3. E. infelix, male, Laurel Co., Ken- 
tucky. 4. E. glenni, female, Adams Co., Ohio. 5. E. tripartitana, female, Morton Co., Kansas. 6. E. infelix, female, Laurel Co., Kentucky. 


Remarks. In describing the forewing of Paedisca 
tripartitana, Zeller (1875) indicated that the interfas- 
cial spot extends from the dorsal margin to the costa 
and is unmarked except for about six very short diago- 
nal lines along the costa. The lines to which he refers 
are dashes of ground color delimiting the costal strigu- 
lae. He also noted that the dorsal edge of the spot is 
considerably wider than the costal edge and that the 
spot'’s distal margin is convex. 

Heinrich (1923) misidentified as E. tripartitana 
some darkly marked Florida specimens of E. scudderi- 
ana, and his illustration of tripartitana male genitalia 
(Fig. 270) is based on one of those specimens. Appar- 
ently he also was referring to the Florida specimens 
when he wrote the diagnostic couplet (no. 11) for tri- 
partitana in his key (p. 137). There he characterized 
the interfascial spot as “extending nearly to costa,” 
which is descriptive of the Florida specimens but is 
not accurate for tripartitana. 

Epiblema tripartitana has been recorded from 
Florida, but the account in Kimball (1965) must be 
viewed with caution. My attempts to match his data 
with museum specimens always resulted in the darkly 
marked form of scudderiana. 


I examined material from Colorado, Connecticut, 
Florida, Illinois, Iowa, Kansas, Louisiana, Michigan, 
Mississippi, Missouri, New Mexico, New York, Ohio, 
and Texas. Forewing length varied considerably: 6 
4.0-7.5 mm (mean = 5.7, n = 17), 9 5.5-10.0 mm 
(mean = 6.7, n = 14). The upper extremes corre- 
sponded to individuals from east Texas reared by Bot- 
timer (1926). The maculation of E. tripartitana ex- 
hibits a certain amount of geographic variation. The 
terminal portion of the forewing usually appears dull 
brown to blackish brown. However, in a small sample 
from Colorado, Iowa, Kansas, and New Mexico, the 
apical area was strongly suffused with light gray, the 
ocellus was poorly defined, and the area above the tor- 
nal angle was conspicuously white. 

Biology. Epiblema tripartitana flies from early 
April to early August, the April and May records being 
from New Mexico or states bordering the Gulf of Mex- 
ico. Bottimer (1926) reported Rudbeckia maxima Nut- 
tall as a laval host in east Texas. He observed early lar- 
val development in the central part of the flower head, 
overwintering of the larva and subsequent pupation in 
the stem at the base of the plant, and spring emer- 
gence of the adult. 


VOLUME 56, NUMBER 4 


279 


| 10 


Fics. 7-10. E. scudderiana. 7. Male, Erie Co., Ohio. 8. Female, Morton Co., Kansas. 9. Male, Bossier Parish, Louisiana. 10. Female, 


Manatee Co., Florida. 


Epiblema glenni D. J. Wright, new species 
(Figs. 1, 4, 11, 15) 


Epiblema infelix (not Heinrich 1923) Miller 1987:58 

(part) (wings, d genitalia). 

Description. Head: Frons white, scales short and 
closely appressed; vertex dark brown, often shading to 
tan anteriorly; outer and ventral surfaces of labial pal- 
pus brown, inner and dorsal surfaces white to tan, apex 
of third segment tan; antenna dark brown above, 
edged with light tan posteriorly, ventral surface pubes- 
cent; ventral surface of scape with varying amounts of 
white to tan scaling. Thorax: Dorsal surface blackish 
brown, ventral surface light tan to dull white; legs dark 


brown outwardly, light tan inwardly, with light tan 
tarsal annulations. Forewing (Figs. 1, 4): ¢ length 
5.0-8.5 mm (mean = 6.1, n = 37), ° length 5.6-8.6 mm 
(mean = 6.6, n = 7); costa nearly straight in males, 
weakly convex basally in females; termen straight to 
weakly concave from M, to R.; male costal fold ex- 
tending from base to approximately 0.6 x length of 
forewing. Upper surface with salmon colored interfas- 
cial spot on dorsal margin, in males extending forward 
to just short of costal fold and separated therefrom by 
narrow strip of dark gray scales, in females extending 
to costa and merging there with whitish to salmon col- 
ored costal strigulae; interfascial spot roughly rectan- 
gular from dorsal margin to fold, constricting abruptly 


11 


Fic. 11. Male genitalia, E. glenni, slide DJW 631 (DJW), scale 
bar 0.5 mm. 


above fold to approximately half its dorsal width, its 
basal margin thinly and intermittently lined with 
white, its dorsal margin with one to three small black 
marks, the latter sometimes weakly expressed or ab- 
sent. Central field of ocellus brown to whitish tan, 
crossed longitudinally by four black dashes, capped at 
apical corner with a black spot of variable expression, 
strongly bordered with gray along basal and tornal 
margins, much more weakly so along distal margin, 
gray border overlaid to varying degrees with light- 
brown to pale-salmon scales. Basal patch blackish 
brown, variously overlaid with gray; male costal fold 
blackish brown with dark gray costal strigulae; median 
area with black spot on basal margin of ocellus just 
above fold, a small gray spot above fold on distal mar- 
gin of interfascial spot, and otherwise marked with 
varying black, brown and gray blotches. Costal area 
beyond interfascial spot brown to blackish brown, 
crossed by four paired white costal strigulae and their 
associated gray stria, the latter running outwardly to- 
ward ocellus and termen; terminal edge of membrane 
lined with white-tipped blackish-gray scales and sepa- 
rated from distal margin of ocellus by narrow strip of 
brown to orange-brown scales; fringe dark blackish 
gray apically, lighter and sometimes brownish at tor- 
nus. Hindwing: Uniformly grayish brown; fringe a 


JRE, MR: 
scale bar 0.5 mm. 


Male genitalia, E. tripartitana, slide D]JW 645 (MEM), 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


————*_ wa Be 


13 


Fic. 13. Male genitalia, E. infelix, slide DJW 755 (USNM), 
scale bar 0.5 mm. 


shade lighter. Male genitalia (Fig. 11): Uncus convex 
and dorsally setose, supported laterally by moderately 
developed shoulders; socii fingerlike, flattened, mod- 
erately long, and covered with long hairlike setae; 
gnathos a narrow sclerotized band, expanded medially; 
aedeagus with approximately 25 deciduous cornuti; 
juxta triangular; caulis short; costal margin of valva 
concave, outer margin weakly convex, inner and outer 
margins of apical one-third of cucullus nearly parallel, 
apex truncated, with rounded corners, invagination of 
ventral margin narrow, width of neck approximately 
one-half that of sacculus, ventral angle acutely 
rounded; margin of basal opening of sacculus featuring 
prominent basal projection with rounded vertex and 
nearly perpendicular sides; clasper mounted on mar- 
gin of basal opening above projection; ventral margin 
of sacculus sparsely setose basally, more densely and 
heavily setose at neck; cucullus densely covered with 
stout setae on distal one-half of inner surface from 
ventral angle to three-fourths distance to apex, outer 
line of setae following distal margin of cucullus to one- 
half distance to apex and migrating thereafter onto in- 
ner surface; apical one-fourth of cucullus densely cov- 
ered with hairlike setae. Female genitalia (Fig. 15): 
Papillae anales narrow, laterally facing, densely setose; 
tergum VIII sparsely setose; anterior and lateral edges 
of sterigma forming U-shaped collar, lamella postvagi- 
nalis well developed, gradually widening posteriorly, 


Fic. 14. Male genitalia, E. scudderiana, slide DJW 198 
(USNM), scale bar 0.5 mm. 


VOLUME 56, NUMBER 4 


281 


Fic. 15. Female genitalia, E. glenni, slide DJW 644 (DJW), 
scale bar 0.5 mm. 


variably setose at the posterolateral corners, posterior 
margin straight, anterior margin A-shaped; posterior 
margin of sternum VII narrow, roundly invaginated to 
depth of 0.3 x length of sterigma, closely approximate 
to sterigma throughout; ductus bursae uniformly nar- 
row, constricted below ostium, with long sclerotized 
patch opposite and posterior to juncture with ductus 
seminalis; corpus bursae with two signa shaped as in 
Fig. 16, one roughly twice the size of the other, the 
larger arising from ventral surface just above mid- 
bursa, the smaller from dorsal surface near juncture 
with ductus bursae. 

Holotype. 3: OH [Ohio]: Adams Co., 1 mi. S. E. of 
Lynx, August 12, 1998, leg. D. J. Wright; genitalia slide 
DJW 631; deposited in USNM. Type locality: 
38°4737’'N, 83°24’197W. 

Paratypes. ILLINOIS: Putnam Co., 17 June 1955, 
MOG (I 2), 8 July 1965, MOG (1 4), 14 July 1968, 


Fic. 16. Female genitalia, E. tripartitana, sterigma and lateral 
view of corpus bursae, slide DJW 751 (DJW), scale bar 0.5 mm. 
MOG (1 4), 20 July 1968, MOG (1 2), 21 July 1965, 
MOG (1 &; genitalia slide DJW 344). KENTUCKY: 
Laurel Co., DBNF, Powerline cut W side Marsh 
Branch Rd, 25 July 1997, LDG (1 6). MICHIGAN: 
Barry Co., T3N RIOW S22, 6 June 1968, JPD (1 d; 
genitalia slide PB 198), T3N R1OW S3, 12 June 1987, 
GJB (1 d). MISSOURI: Reynolds Co., Grasshopper 
Hollow, Prairie Fen (TNC), 16 September 1996, GJB 
(1 °; genitalia slide DJW 346); St. Francois Co., St. 
Francois St. Pk. along Coonville Creek, 10 July 1982, 
GJB (1 2). OHIO: Adams Co., 1 mi. SE of Lynx, 25 
July 1998, DJW (1 2; genitalia slide DJW 630), 1 Au- 
gust 1997, DJW (1 ¢; genitalia slide DJW 643), 3 Au- 
gust 1998, DJW (2 4), 3 August 2000, DJW (5 2), 12 
August 1998, (7 d, 2 2; d genitalia slides DJW 458, 632, 
? genitalia slides DJW 457, 644), 21 August 1993, 
DJW (1 ¢; genitalia slide DJW 323), 10 September 
1988, DJW (1 ¢; genitalia slide DJW 172), Lynx Prairie 


#2, 10 September 1988, LDG (1 ¢; genitalia slide 
LDG 87); Erie Co., Resthaven Wildlife Area, 13 July 
1991, LDG (1 4); Greene Co., Wright-Patterson Air 
Force Base, Huffman Prairie, Site G-3, 24 July 1992, 
EHM (1 ¢); Hamilton Co., Cincinnati, 3349 Morrison 
Ave., 24 July 1997, DJW (1 2); Scioto Co., Brushy Fork 
near Upper Twin Creek, 26 May 1990, DJW (1 d; gen- 
italia slide DJW 347). Paratype depositories: AMNH, 
GJB, UConn, LDG, INHS, UL, MSU, MEM, MCZ, 
OL, USNM, DJW. 

Etymology. This species is named in honor of the 
late Murray O. Glenn, in recognition of his contribu- 
tion to the knowledge of the microlepidoptera of cen- 
tral Illinois. 

Distribution and biology. The sample of 78 spec- 
imens from Illinois, Kentucky, Michigan, Missouri, 
North Carolina, Ohio, and Tennessee indicates a flight 
period from late May to mid September, but three- 
fourths of those records occurred between mid July 
and mid September. No larval host has been recorded. 

Diagnosis. Epiblema glenni differs from E. tripar- 
tiana in the shape and coloration of the interfascial 
spot. In E. tripartitana the spot appears white to the 
naked eye, but a pale salmon tint usually is detectable 
under magnification. In females the spot merges with 
the costal stigulae, forming a continuous band from 
dorsal margin to costa; in males it is intercepted by the 
costal fold, but light-gray to gray costal strigulae on the 
adjacent portion of the fold often create the impres- 
sion that it continues to the costal margin. The convex 
curvature of its distal margin varies from circular to 
that of a line bent just above the fold. In E. glenni the 
interfascial spot is distinctly salmon colored. Its width 
narrows markedly above the fold, where the distal 
edge angles abruptly inward. In females it continues 
forward to the costa as a narrower band: in males it is 
separated from the costal fold by a narrow strip of dark 
scales. I was unable to find characters of the male or 
female genitalia that would separate the two species. 
Although Figs. 11 and 12 suggest subtle differences in 
the shape of the cucullus, this feature is variable in 
both species and unreliable for diagnostic purposes. 


Epiblema infelix Heinrich 
(Figs. 3, 6, 13, 17) 


Epiblema infelix Heinrich 1923:151, Fig. 276 (genitalia 
of ¢ holotype); McDunnough 1939:48: Powell 
1983:35; Miller 1987:58 (part) (2 genitalia). 
Holotype. ¢: Fiske Coll., 25 May 1904, Tryon, N.C.; 

USNM Type No. 24828; genitalia slide USNM 72880. 
Paratypes. NORTH CAROLINA: Tryon, 4 July 

1904, Fiske (1 2; genitalia slide USNM 70802), 5 July 

1904, Fiske (1 d; genitalia slide USNM 70801). 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 17. Female genitalia, E. infelix, left apophysis anterior 
omitted for clarity, slide DJW 677 (MEM), scale bar 0.5 mm. 

Remarks. Epiblema infelix has been a source of 
confusion for some time. It is poorly represented in in- 
stitutional collections, and I often found mixed series 
in the material that I borrowed. I examined the Michi- 
gan specimen pictured in Miller (1987:58) as E. infelix 
and found it to be E. glenni. Miller's drawing of the 
male genitalia is representative of glenni, but his illus- 
tration of the female sterigma is base on the paratype 
of infelix from North Carolina (Miller pers. com.). 


VOLUME 56, NUMBER 4 


Heinrich’s description portrays the forewing color as 
brownish ochreous and the white interfascial spot as 
irregularly square, with one or two short fuscous 
dashes on the dorsal margin. Based on an examination 
of 45 specimens, including the holotype and both 
paratypes, I would characterize the predominant hue 
of the forewing as gray, but worn specimens tend to 
have a brownish appearance. The basal area is rather 
dark, appearing blackish gray to the naked eye. The 
white interfascial spot is roughly triangular, extending 
from the dorsal margin forward to approximately two- 
thirds the distance to the costa. Its sharply defined 
basal edge angles obliquely outward from the dorsal 
margin and often contains a small, basally directed, tri- 
angular projection just forward of the dorsal margin. 
The distal edge is less well defined, the outer vertex on 
the dorsal margin being irregularly truncated. The in- 
terfascial spot is invaded from the dorsal margin by 
one to three dark marks, which vary considerably in 
degree of expression. Orange-brown coloration occurs 
in the costal area beyond the interfacial spot and along 
the distal edge of the ocellus, the former area being 
crossed by four paired costal strigulae and their associ- 
ated gray stria. Forewing length varied as follows: 6 
6.2-8.7 mm (mean = 7.6, n = 20), 9 9.5-10.2 mm 
(mean = 9.8, n = 4). The male costal fold extends from 
base to approximately 0.5 x length of forewing. 

Distribution and biology. I studied specimens 
from Alabama, Arkansas, Kentucky, Michigan, Missis- 
sippi, North Carolina, South Carolina, and Virginia. 
Except for one North Carolina record dated 22 Octo- 
ber, they indicate a flight period from early April to 
early July. Eighty percent of the records occurred be- 
tween mid April and mid June. No larval host has been 
recorded. 


Epiblema scudderiana (Clemens) 
(Figs. 7-10, 14, 18) 


Hedya scudderiana Clemens 1860:358. 

Euryptychia saligneana Clemens 1865:141. 

Paedisca affusana Zeller 1875:307. 

Eucosma scudderiana Fernald {1903]:459; Barnes and 
McDunnough 1917:171. 

Epiblema scudderiana Heinrich 1923:147; McDun- 
nough 1939:48; Powell 1983:35; Miller 1987:57 
(wings, d and ° genitalia). 

Lectotype. Hedya scudderiana, designated by Dar- 
lington (1947:95). d: Type 7215, Paedisca scudderiana 
Clem.; Academy of Natural Sciences of Philadelphia. 
Type locality: Massachusetts. Miller (1973: Fig. 40) 
provides a photograph of this specimen and a discus- 
sion of the type fixation difficulties associated with 
Clemens’ species. 


af 
err 


Fic. 18. Female genitalia, E. scudderiana, left apophysis ante- 
rior omitted for clarity, slide DJW 647 (INHS), scale bar 0.5 mm. 


Remarks. The basic forewing pattern of E. scudde- 
riana has the following features: basal patch blackish 
brown, variably overlaid with blue-gray scales; inter- 
fascial spot white, often marked with varying amounts 
of blue-gray scaling, extending from dorsal margin to 
just short of costa, its basal and distal margins roughly 
convex, producing a bulletlike shape; ocellus crossed 


longitudinally by up to four black dashes and variably 


bordered on basal, distal and tonal margins with blue- 
gray scales; a narrow band of orange-brown coloration 
along costa beyond interfascial spot, crossed by four 
paired white costal strigulae, the latter with associated 
blue-gray stria running distally toward ocellus and ter- 
men; median area with prominent black spot just 
above fold on basal edge of ocellus and a variably ex- 
pressed blue-gray band along distal margin of interfas- 
cial spot. There is considerable variation in the amount 
of contrast between interfascial spot and median area 
and in the coloration of the ocellus. In the Florida 
population that Heinrich mistook for tripartitana, the 
interfascial spot is nearly immaculate, the median area 
is dark brown, with little if any white scaling, and the 
central field of the ocellus is tan to brown. At the other 
extreme, specimens from the upper Midwest feature 
an interfascial spot that is moderately speckled with 
blue gray, a median area that is covered predominantly 
with white and bluish-gray scales, creating a mottled 
pale gray appearance to the naked eye, and an ocellus 
with a white central field. The study sample of 93 spec- 
imens included numerous intermediates from various 
locations, including Florida, New England, and the 
Gulf coast. 

I examined male and female genitalia preparations 
of specimens representative of the range of the insect 
and found the following characters most useful for di- 
agnostic purposes. Males (n = 14): Uncus semicircular, 
dorsally setose, supported laterally by well developed 
shoulders; socii short, fingerlike, flattened, and moder- 
ately setose; costal margin of valva concave, outer mar- 
gin convex, invagination of ventral margin broad and 
shallow, ventral angle gently rounded; margin of basal 
opening of sacculus strongly sinuate; inner surface of 
cucullus sparsely covered on basal one-third with hair- 
like setae, densely so on distal two-thirds with stouter 
setae. Females (n = 17): Papillae anales narrow, later- 
ally facing and strongly setose; anterior and lateral 
margins of sterigma forming U-shaped collar, its paral- 
lel sides flaring posteriorly; lamella postvaginalis well 
developed, with semicircular anterior margin and up 
to five setae on each lateral margin; posterior margin 
of sternum VII broad, concavely invaginated to depth 
of 0.5 x length of sterigma, approximate to ostium me- 
dially, diverging from sterigma laterally; ductus bursae 
tapering gradually, constricted just below ostium, and 
mildly schlerotized opposite juncture with ductus sem- 
inalis; corpus bursae with two cone-shaped signa of 
nearly equal size, positioned opposite one another at 
roughly mid-bursa. Figures 14 and 18 are typical of the 
dark population from Florida. Other genitalia illustra- 
tions can be found in Heinrich (1923, Figs. 270 (la- 
beled tripartitana) and 271) and Miller (1987:57). 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Distribution and biology. Epiblema scudderiana 
occurs across the eastern half of the United States and 
southern Canada. I examined specimens from Al- 
abama, Florida, Georgia, Iowa, Kansas, Louisiana, 
Massachusetts, Mississippi, New York, North Carolina, 
Ohio, and Vermont. The larva is known to be a late- 
instar stem feeder on various species of Solidago, 
where it produces a conspicuous gall. Miller (1976) re- 
ports on the biology of this insect, providing additional 
larval hosts and references to the literature on this 
subject. 


DISCUSSION 


Of the four species treated here, specimens in rea- 
sonably good condition can be separated by features of 
the interfascial spot. In E. scudderiana it is white and 
stops just short of the costa. Its large bulletlike shape is 
distinctive, but in lighter colored specimens its distal 
edge is often poorly defined. Epiblema infelix has a 
white, roughly triangular interfascial spot, the anterior 
vertex of which extends about two-thirds the distance 
from the dorsal margin to the costa. The color and 
shape of the spot in E. tripartitana and E. glenni are 
discussed in the diagnosis section under glenni. 

The dark form of E. scudderiana from Florida has a 
prominent blue-gray band that extends nearly from 
the costa to the dorsal margin along the distal edge of 
the interfascial spot. In specimens from the upper 
Midwest this band is not so well defined and is often 
broken by white coloration protruding from the inter- 
fascial spot. Intermediate expressions of this band 
were found in material from various localities. In E. 
tripartitana and E. glenni this feature is reduced to a 
short blue-gray dash above the fold and, in some in- 
stances, another disjoint blue-gray spot nearer the 
costa. Epiblema infelix exhibits varying amounts of 
gray shading beyond the interfascial spot, but the con- 
trast with neighboring ground color is slight at best. 

Epiblema glenni and E. tripartitana, though similar 
in both male and female genitalia, are distinct in this 
respect from infelix and scudderiana, as are the latter 
two from each other. Differences in valval shape are il- 
lustrated in Figs. 11-14. The margin of the basal open- 
ing of the sacculus in glenni and tripartitana features a 
basal projection below the clasper with rounded apex 
and sides that are roughly perpendicular. Epiblema in- 
felix has no such projection, and the margin in scudde- 
riana is strongly sinuate. The shape, relative size, and 
position of the female signa are useful diagnostic char- 
acters. In glenni and tripartitana the signa are unequal 
in size (Fig. 16), the larger arising near mid-bursa, the 
smaller near juncture with ductus bursae. In infelix 
and scudderiana they are roughly equal in size and 


VOLUME 56, NUMBER 4 


arise opposite one another at approximately mid-bursa 
(Figs. 17, 18). 


ACKNOWLEDGMENTS 


I thank R. L. Brown for advice and encouragement, D. L. Wag- 
ner for photographing the holotype of E. tripartitana, E. H. Metzler 
for help with the adult photographs, J. Snider for the use of his lab- 
oratory, and W. E. Miller and an anonymous reviewer for helpful 
comments on the manuscript. I especially appreciate the confidence 
and cooperation of Paul Knoop, whose permission to collect on his 
property in Adams, Co., Ohio, led to the results in this paper. 


LITERATURE CITED 


BARNES, W. & J. McDuNNoucH. 1917. Checklist of the Lepidoptera 
of boreal America. Herald Press, Decatur, Illinois. 392 pp. 

BOTTIMER, L. J. 1926. Notes on some Lepidoptera from eastern 
Texas. J. Agr. Research 33:797-819. 

Brown, R. L. & J. A. POWELL. 1991. Description of a new species 
of Epiblema (Lepidoptera: Tortricidae: Olethreutinae) from 
coastal redwood forests in California with an analysis of 
forewing pattern. Pan-Pac. Entomol. 67:107—114. 

CLEMENS, B. 1860. Contributions to American lepidopterology. 
No. 6. Proc. Acad. Nat. Sci. Philadelphia 1860:345-362. 

. 1865. North American micro-lepidoptera. Proc. Entomol. 
Soe. Philadelphia 5:133-147. 

FERNALD, C. H. [1903]. In Dyar, H. G., A list of North American 
Lepidoptera. U.S. Nat. Mus. Bull. 52:1-723. 


285 


DaRLINGTON, E. P. 1947. Notes on certain types of Lepidoptera 
described by Brackenridge Clemens. Trans. Am. Entomol. Soc. 
73:85-104. 

HEINRICH, C. 1923. Revision of the North American moths of the 
subfamily Eucosminae of the family Olethreutidae. U.S. Nat. 
Mus. Bull. 123:1-298. 

KIMBALL, C. P. 1965. Arthropods of Florida and neighboring land 
areas. Vol. 1. Florida Dept. of Agriculture. 363 pp. 

McDunnoucu, J. 1939. Check list of the Lepidoptera of Canada 
and the United States of America. Part II. Microlepidoptera. 
Mem. South. Calif. Acad. Sci. 2:3-171. 

MILLER, W. E. 1973. Clemens types of Olethreutinae (Lepi- 
doptera, Tortricidae). Trans. Am. Entomol. Soc. 99:205-234. 

. 1976. Biology and taxonomy of three gall forming species 

of Epiblema (Olethreutidae). J. Lepid. Soc. 30:50-58. 

. 1987. Guide to the olethreutine moths of midland North 
America (Tortricidae). U.S.D.A. For. Serv. Agric. Handbook 
660:1—-104. 

POWELL, J. A. 1983. Tortricidae, pp. 31-41. In Hodges, R. W. et al. 
(eds.), Check list of the Lepidoptera of America north of Mex- 
ico. E. W. Classey & Wedge Entomol. Res. Foundation, Lon- 
don, England. 

ZELLER, P. C. 1875. Beitriige zur Kenntniss der nordamericanis- 
chen Nachtfalter, besonders der Microlepidopteren. Verh. 
Zool.-bot. Ges. Wien. 25:205-360, pl. VITI-X. 


Received for publication 30 January 2002; revised and accepted 2 
June 2002. 


GENERAL NOTES 


Journal of the Lepidopterists’ Society 
56(4), 2002, 286-288 


IMMATURE STAGES OF ETEONA TISIPHONE (NYMPHALIDAE: SATYRINAE) 


Additional key words: 


Immatures of most Neotropical Satyrinae are still 
poorly known or undescribed. The available informa- 
tion (including some partial or incomplete descrip- 
tions) includes species of rather few genera that repre- 
sent only a small fraction of the main sections of this 
subfamily (see Singer et al. 1971, Kendall 1978, Singer 
et al. 1983, Young 1984, DeVries 1987, Sourakov 1996, 
Pelz 1997). The present paper describes the immature 
stages of Eteona tisiphone (Boisduval, 1836), a sub- 
montane Pronophiliti, in comparison with other 
known Satyrinae immatures. 

Study sites and methods. Adults and immatures 
of Eteona tisiphone were studied in the field in five 
different localities in SE Brazil: Alto da Serra, 
Paranapiacaba, Sao Paulo (1000-1100 m); Serra do 
Japi, Jundiai, Sao Paulo (1100-1200 m); Santa Gene- 
bra Forest Reserve, Campinas, Sao Paulo (600-650 
m); Campos do Jordao State Park, Campos do Jordao, 
Sao Paulo (1500-2000 m); and Serra do Cristo, Pogos 
de Caldas, Minas Gerais (1400 m). Immatures were 
collected in the field; eggs were also obtained from 
fertile females confined in a plastic bag with leaves of 
the host plant. Larvae were reared in plastic contain- 
ers cleaned daily, with fresh plant material provided 
every two or three days (following Freitas 1991). Data 
were taken on behavior and development times for all 
stages, and head capsules and pupal castings were 
preserved. When there was sufficient material, imma- 
tures were fixed in Kahle (AVLF collection). Nomen- 
clature follows Miller (1968) modified by Harvey 
(1991), who treated the group as a subfamily, and 
downranked Miller’s subfamilies and tribes to tribes 
and subtribes, respectively. 

Host plants oviposition, and immature behav- 
iors. The host plants of Eteona tistphone are bamboo 
species in the genus Chusquea (in mountain sites) and 
Merostachys (only in Campinas). Oviposition was ob- 
served many times in all study sites. 

Females usually lay eggs near midday, when the 
temperature is high. Eggs are laid singly on the 
shoot tips of growing stems of bamboos, indepen- 
dent of their height above the ground (observed 
from 1 to6m high). Eggs are usually laid on young 
branches without leaves: the larva srouldl then feed 

on the newly sprouting young leaves. The female 
lands on the tip of the stem and touches the tip of 


bamboo feeders, Pronophiliti, Zetheriti. 


her abdomen near a bamboo node (with or without 
new leaves) until she finds a suitable place to lay an 
egg. This process is repeated another 4 to 6 times, 
after which the female rests for some time before 
laying additional eggs or moving to another bamboo 
patch. 

The young larva eats part of the egg chorion, and af- 
ter some time starts to feed on the new bamboo leaves. 
The larva rests along a bamboo leaf with the head usu- 
ally towards the leaf apex, being perfectly hidden and 
difficult to locate in the field. The larva is very active 
when touched and usually wanders a lot when con- 
fined in small dishes. 

Description of early stages. The following de- 
scriptions are based on material reared from the Serra 
do Japi. All features of the immatures are very similar 
in all additional locations. 

Egg (Fig. la). Spherical, slightly truncated at base, 
with very fine reticulations (similar to minute pits 
when subjected to cross-lighting); light yellow, becom- 
ing more reddish with a dark cap (head capsule of 
growing larva) near eclosion. Height 1.26 mm, diame- 
ter 1.14 mm. Duration: 6 days (n = 6). 

First instar (Figs. 1b, 2). Head capsule dark brown, 
with a pair of short scoli on vertex, and six pairs of reg- 
ularly spaced spatulate setae. Head capsule width 
0.80—0.86 mm (n = 4): head scoli 0.14—0.12 mm (n = 
4). Body beige (light green after feeding), smooth, 
with many poorly visible red longitudinal stripes, and a 
pair of short caudal filaments. Body chaetotaxy illus- 
trated in Fig. 2. Maximum length 7 mm. Duration: 
4-5 days. 

Second instar. Head dark brown with light frons 
and two dark brown diverging scoli on vertex. Head 
capsule width 1.02-1.08 mm (n = 3); scoli 0.84—0.90 
min (n = 3). Body slender, light green with many lon- 
gitudinal white stripes; caudal projections short. Maxi- 
mum length 12 mm. Duration: 4-6 days. 

Third instar (Fig. 1c). Head the same as in previ- 
ous instar. Head capsule width 1.36—1.50 mm (n = 4); 
scoli 1.60-1.80 mm (n = 4). Body dark green with a 
broad lateral longitudinal white stripe and many addi- 
tional narrow light stripes; caudal projections medium 
long (similar to head scoli). Maximum length 16 mm. 
Duration: 4-5 days. 

Fourth instar (Fig. 1d). Very similar to third instar, 


VOLUME 56, NUMBER 4 


287 


Fic. 1. Early stages of Eteona tisiphone. a, egg; b, first instar; , third instar; d, fourth instar; e, f, fifth instar; g, h, pupa (ventral, lateral). 


with patterns in general darker than in previous in- 
stars. Head capsule width 1.80-2.04 mm (n = 4); scoli 
2.34-3.00 mm (n = 4). Maximum length 27 mm. Du- 
ration: 4—5 days. 

Fifth instar (Fig. le-f). Head light brown with lat- 
eral area dark; a pair of long normally slightly diverg- 
ing scoli (convergent in one of two individuals from 
Paranapiacaba). Head capsule width 2.73-2.77 mm (n 
= 3); scoli 3.74-3.90 mm (n = 3). Body slender, dark 
green; a conspicuous lateral longitudinal white stripe 
broadening towards the abdomen and many additional 


narrow dark stripes; caudal projections medium long. 
Maximum length 40 mm. Duration: 9-10 days. 

Pupa (Fig. 1g-h). Elongated, smooth, with long 
pointed ocular caps; light green with many whitish ar- 
eas. Total length 25 mm. Duration 9 days. 

Discussion. Although it is widely accepted that 
Eteona is a genus belonging to the Neotropical sub- 
tribe Pronophiliti (Miller 1968, Brown 1992), its im- 
matures diverge greatly from all known Pronophiliti, 
whose larvae have short diverging horns and a short bi- 
fid tail, and whose pupae are short and stubby with 


Fic. 2. Chaetotaxy (upper) and head capsule (front view) 
(lower) of the first instar larva of Eteona tisiphone. 


short ocular caps (DeVries 1987, Sourakov 1996, Pelz 
1997). In contrast to other Pronophiliti, the larvae of 
Eteona have a pair of long horns and a medium bifid 
tail, and the pupa is slender with long ocular caps. This 
makes Eteona different from most Neotropical Satyri- 
nae (Singer et al. 1983, DeVries 1987, AVLF unpub- 
lished data from 59 species), except perhaps some 
Splendeuptychia (also a bamboo feeder) that also have 
elongated larvae and pupae (AVLF unpublished). 
Comparison of these immatures with the described 
immatures of Zethera (Wolfe 1996), Penthema, and 
Neorina (Elymniini, Zetheriti) (Lee & Chang 1988, Li 
& Zhu 1992, Lee & Wang 1995, Wolfe 1996) shows 
surprising similarities (especially in the individuals 
with converging head scoli), except in the first instar 
head capsule (smooth and without scoli in the known 
Zetheriti). This similarity could be due to camouflage 
among bamboo leaves, a feature present also in the 
Zetheriti and also in Splendeuptychia. However, the 
simple fact of being a bamboo feeder does not neces- 
sarily lead to an elongated larva and pupa, as con- 
firmed by the rearing of other species of Splendeupty- 
chia, and also Forsterinaria, Carminda, Moneuptychia 
and Pedalioides (AVLF unpublished). 

In sum, the immatures of E. tistphone are highly 
divergent from other known Pronophiliti and sug- 
gest that the taxonomic position of this genus should 
be verified. Any additional information from early 
stages of Pronophiliti will be valuable to help to un- 
derstand this group and the relationships within the 
Satyrinae 

This study was conducted as part of a Post Doctoral 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


project on Satyrinae biology and systematics (BIOTA- 
FAPESP program, grants 98/05101-8 and 00/01484-1). 
I would like to thank Dr. Keith S. Brown Jr. for reading 
the manuscript, and the Fundagéo José Pedro de 
Oliveira for allowing fieldwork in the Reserva Mata da 
Santa Genebra. | 


LITERATURE CITED 


Brown, K. S. Jr. 1992. Borboletas da Serra do Japi: Diversidade, 
habitats, recursos alimentares e variagao temporal, pp. 142-187, 
18 figs. In Morellato, L. P. C. (ed.), Hist6ria natural da Serra do 
Japi. Ecologia e preservacao de uma area florestal no sudeste do 
Brasil. Campinas, Editora da Unicamp/Fapesp. 

DEVRIES, P. J. 1987. The butterflies of Costa Rica and their natural 
history. Papilionidae, Pieridae, Nymphalidae. Princeton Uni- 
versity Press, Princeton, New Jersey. 

DEVRIES, P. J., I. J. KircHiNc & R. I. VANE-WRriIGHT. 1985. The sys- 
tematic position of Antirrhea and Caerois, with comments on 
the higher classification of the Nymphalidae (Lepidoptera). 
Syst. Entomol. 10:11-32. 

Freitas, A. V. L. 1991. Variagao morfolégica, ciclo de vida e sis- 
tematica de Tegosa claudina (Eschscholtz) (Lepidoptera, 
Nymphalidae, Melitaeinae) no Estado de Sao Paulo, Brasil. 
Rey. bras. Ent. 35:301-306. 

Harvey, D. J. 1991. Higher classification of the Nymphalidae 
(Appendix B), pp. 255-273. In Nijhout, H. F., The develop- 
ment and evolution of butterfly wing patterns, Smithsonian 
Press. 

KENDALL, R. O. 1978. Larval foodplant, life history notes and tem- 
poral distribution for Splendeuptychia kendalli (Satyridae) from 
Mexico. J. Lepid. Soc. 32:86-87. 

LEE, J.-Y. & Y.-C. CHANG. 1988. The illustrations of butterflies in 
Taiwan. Taipei: Taiwan Mus. pp. 72-73. 

LEE, J.-Y. & H.-Y. WANG. 1995. The illustrations of butterflies on 
Kinmen and Matsu islands. Taipei: Taiwan Mus. pp. 145-147. 

Li, C. L. & B. Y. ZHu. 1992. Atlas of Chinese butterflies. Shanghai: 
Shanghai Yuandong. 238 pp. 

MILLER, L. D. 1968. The higher classification, phylogeny and zoo- 
geography of the Satyridae (Lepidoptera). Mem. Am. Entomol. 
Soc. 24:iii + 174 pp. 

PELZ, V. 1997. Life history of Pedalioides parepa from Ecuador 
(Lepidoptera: Nymphalidae: Satyrinae). Trop. Lepid. 
8(1):41-45. 

SINGER, M. C., P. J. DEVrRIES & P. R. EHRLICH. 1983. The Cissia 
confusa species-group in Costa Rica and Trinidad (Lepidoptera: 
Satyrinae). Zool. J. Linn. Soc. 79:101-119. 

SINGER, M. C., P. R. EHRLICH & L. E. GILBERT. 1971. Butterfly 
feeding on Lycopsid. Science 172:1341-1342. 

Sourakoy, A. 1996. Notes on the genus Calisto, with descriptions 
of the immature stages (Part I) (Lepidoptera: Nymphalidae: 
Satyrinae). Trop. Lepid. 7:91-112. 

WOLFE, K. V. 1996. Notes on the early stages of Zethera musides 
(Lepidoptera: Nymphalidae: Satyrinae). Trop. Lepid. 
7:147-150. 

Younc, A. M. 1984. Natural history notes for Taygetis andromeda 
(Cramer) (Satyridae) in Eastern Costa Rica. J. Lepid. Soe. 
38:102-113. 


ANDRE V. L. FREITAS, Museu de Histéria Natural 
and Departamento de Zoologia, Instituto de Biologia, 
Universidade Estadual de Campinas, CP 6109, 
Campinas, Séo Paulo, 13083-970, Brazil 


Received for publication 7 February 2002; revised and accepted 30 
May 2002. 


Journal of the Lepidopterists’ Society 
56(4), 2002, 289-290 


NOTES ON THE LARVAL DIET OF THE PAINTED LICHEN MOTH HYPOPREPIA FUCOSA HUBNER 
(ARCTIIDAE: LITHOSIINAE) 


Additional key words: Trentepohlia, Cladonia polycarpoides, Physcia millegrana, algal partners, algivory. 


It is well known that the caterpillars of the subfam- 
ily Lithosiinae (Arctiidae), or lichen moths, feed on 
lichens. They are suspected of being primarily algi- 
vores, feeding on algae, either free-living, or as a 
lichen symbiont (T. McCabe pers. com., Rawlins 
1984 and pers. com.). For most species, even the 
most basic information on the larval diet is lacking or 
is poorly documented (N. Jacobson pers. com., P. 
Opler pers. com., J. Rawlins 1984 and pers. com.). 
Reports of food sources for this subfamily consist pri- 
marily of generalizations such as “lichens,” “algae,” 
“mosses,” or the plants that the larvae were observed 
on (Tietz 1952, 1972, Forbes 1954, Covell 1984, 
Rawlins 1984, McCabe 1991, Wagner et al. 1997, 
Robinson et al. 2001, D. Schweitzer pers. com.). 
Identification of host taxa species, or even to genus, is 
rarely reported. In this note we describe our observa- 
tions of feeding by Hypoprepia fucosa Hiibner and 
identify an algal host and two lichens ingested in cap- 
tivity. We also identify the algal partners of the 
lichens. We are aware of only one other unpublished 
account that identifies a specific lichen or algal host 
for this species: a report by T. McCabe (pers. com.) 
of H. fucosa feeding on the free-living algae Proto- 
coccus viridis C. A. Agardh. 

On 10 July 2001 we observed H. fucosa feeding on 
a free-living alga in the genus Trentepohlia. The 
species could not be determined with confidence, but 
may have been T. aurea Mart. given the strong orange 
pigmentation (P. DePriest pers. com.). The alga was 
growing on a concrete fireplace at a campsite located 
in a mature mixed-oak forest, Plumstead Township, 
Bucks County, Pennsylvania. The forest canopy con- 
sisted of white oak (Quercus alba L.), red oak (Quer- 
cus rubra L.), sugar maple (Acer saccharum Marsh.) 
and chestnut oak (Quercus prinus L.). The caterpillar 
was observed feeding on the algae for about one hour 
beginning at 2130 h before it and the alga was col- 
lected for subsequent identification. 

The caterpillar was first observed about one hour 
after torrential rainfall associated with strong thun- 
derstorms had passed through the area. The temper- 
ature was near 24°C and the air was humid. The 
caterpillar was contained within a small glass jar with 
no available food source for approximately 24 hours, 
then transferred to a small plastic insect cage lined 
with a 0.4 cm deep layer of garden soil. The caterpil- 
lar was then supplied with two lichens obtained from 


the trunks of a red maple (Acer rubrum L.) and a red 
oak from a suburban yard in East Brunswick, Mid- 
dlesex County, New Jersey. The caterpillar readily ac- 
cepted these food sources and fed on both intermit- 
tently for the next three days before succumbing to a 
braconid wasp parasite (D. Wagner pers. com.). No 
preference for either lichen was apparent. The lichen 
collected on the red maple was Cladonia polycar- 
poides Nyl. and the lichen from the red oak was 
Physcia millegrana Degel. Algal partners of C. poly- 
carpoides are from the Trebouxia irregularis (Hil- 
dreth et Ahmadjian) group (T. irregularis, T. glomer- 
ata, T. pyriformis, T. excentrica, T. magna), and for P. 
millegrana, from the Trebouxia impressa (Ahmad- 
jian) Archibald group (T. impressa, T. gelatinosa) (P. 
DePriest pers. com.). 

While feeding on the algal host and the lichens in 
captivity, the caterpillar used a scraping motion that 
may have gleaned algae from the substrate or re- 
moved the fungal component to obtain the lichen 
cortex with algae (J. Rawlins pers. com.). Rawlins 
(pers. com.) and McCabe (pers. com.) have sug- 
gested that H. fucosa and many other lithosiines are 
algal feeders, and Rawlins (pers. com.) has success- 
fully reared more than 40 species of lithosiines from 
more than 20 genera worldwide on free algae grow- 
ing on bark, twigs, and leaves of woody plants, often 
intermixed with lichens, small fungi, sooty molds, and 
mosses. Many reports of lithosiines feeding on 
lichens, mosses, or other substrates may be incidental 
to ingestion of algae growing mixed with these mate- 
rials (McCabe 1991 and pers. com., J. Rawlins pers. 
com.), but the relationship between many species 
and algivory requires additional investigation. Our 
observations seem to support an algal larval diet for 
H. fucosa, and we hope add to understanding of 
lithosiine algivory. 


We are indebted to Paula DePriest of the Smithsonian Institu- 
tion for identifying the lichens and algae, to David Wagner of the 
University of Connecticut for assistance identifying H. fucosa and 
to John Rawlins of the Carnegie Museum of Natural History and 
Tim McCabe of the New York State Museum for sharing their un- 
derstanding of the biology and larval diet of H. fucosa and other 
lithosiines. We are grateful to John Rawlins, Paula DePriest, Carla 
Penz and an anonymous reviewer for manuscript review and help- 
ful comments. Thanks are also due to Boy Scouts of America Troop 
220 for the opportunity to observe H. fucosa and to EcolSciences, 
Inc. for the time and financial resources necessary to prepare this 
manuscript. 


LITERATURE CITED 


CovELL, C.V,, Jr. 1984. A field guide to the moths of eastern North 
America. Houghton Mifflin Company. Boston. xv + 496 pp. 
Fores, W. T. M. 1960. The Lepidoptera of New York and neigh- 
boring states: Part IV. Agaristidae through Nymphalidae includ- 
ing butterflies. Memoir 371. Cornell University Agricultural Ex- 

periment Station, Ithaca, New York. 188 pp. 

McCasg, T. 1991. Atlas of Adirondack caterpillars with a host list, 
rearing notes and a selected bibliography of works depicting 
caterpillars. New York State Museum Bulletin No. 470, iv + 
114 pp. 

RAWLINS, J. E. 1984. Mycophagy in Lepidoptera. Chapter 15, pp. 
382-423. In Wheeler, Q. & M. Blackwell (eds.), Fungus-insect 
relationships. Perspectives in ecology and evolution. Columbia 
University Press, New York. 514 pp. 

ROBINSON, G. S., P. R. ACKERY, G. W. BECCALONTI, I. J. KITCHING & 
L. M. HERNANDEZ. 2001. HOSTS—a database of the host- 
plants of the world’s Lepidoptera. Electronic resource. The 
Natural History Museum, London, England. 

Tietz, H. M. 1952. The Lepidoptera of Pennsylvania: a manual. 
The Pennsylvania State College School of Agriculture Ag- 


Journal of the Lepidopterists’ Society 
56(4), 2002, 290-292 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


ricultural Experiment Station. State College, Pennsylvania. 

194 pp. 

. 1972. An index to the described life histories, early stages 
and hosts of the Macrolepidoptera of the continental United 
States and Canada. Vol. I and I. The Allyn Museum of Ento- 
mology, Sarasota, Florida. 1041 pp. 

WAGNER, D.L., V. GILES, R.C. REARDON & M. L. MCManus. 1997. 
Caterpillars of eastern forests. U. S. Dept. of Agriculture Forest 
Service, Forest Health Technology Enterprise Team Publica- 
tion 96-34. 113 pp. 


Davip P. Moskowitz, EcolSciences, Inc. 75 Fleet- 
wood Drive, Suite 250, Rockaway, New Jersey 07866, 
USA dmoskowi@ecolsciences.com, AND  CLAUDIS 
WESTPHAL, 22 Appleby Lane, East Brunswick, New 
Jersey OSS16, USA 


Received for publication 30 January 2002; revised and accepted 17 
June 2002. 


MEGATHYMUS YUCCAE IN KENTUCKY (HESPERIIDAE, MEGATHYMINAE) 


Additional key words: Yucca Giant-Skipper, Yucca filamentosa. 


On 11 April 2001 the third author discovered a 
colony of Yucca Giant-Skippers, Megathymus yuccae 
(Boisduval & Leconte) (Megathymidae), in southeast- 
ern Calloway County, Kentucky, approximately 22 kin 
(13.5 mi) east-south-east of Murray in the western part 
of the state. At least 8—10 individuals were observed, 
two or three of which were females, obvious because of 
their larger overall size and thicker abdomens. Males 
were observed flying about in pursuit of one another— 
apparently defending territories—and perching on the 
ground and vegetation. One male specimen was col- 
lected and photos were taken of several individuals, in- 
cluding a mating pair. A second mated pair also was ob- 
served, and on one occasion three males were observed 
simultaneously in pursuit of a single female. 

The Yucca Giant-Skippers were found amidst a 


a 


; Fic. 1. Yucca Giant-Skipper, Megathymus yuccae, Calloway 
Co., Kentucky, 13 April 2001; photo by Loran D. Gibson. 


Fic 2. Underside of the Yucca Giant-Skipper, Megathymus 
yuccae, Calloway Co., Kentucky, 13 April 2001; photo by Loran D. 
Gibson. 


VOLUME 56, NUMBER 4 


Fic. 3. Yucca Giant-Skipper food plant, Yucca filamentosa L., with larval “chimney,” Calloway Co., Kentucky, 18 April 2002; photo by W. 


R. Black, Jr. 


colony of Yucca, Yucca filamentosa L. (Liliaceae), con- 
sisting of approximately two or three dozen plants scat- 
tered along the shoulders of a gravel road. The site ex- 
tended for approximately 245 m (200 yards) and was 
situated on a ridgetop of dry, gravelly soils. The sur- 
rounding vegetation consisted primarily of relatively 
young, second-growth upland oak-hickory forest. 
There were also numerous Loblolly pines, Pinus taeda 
L. (Pinaceae), a species not native to Kentucky, scat- 
tered throughout the forest and roadside. The road- 
side shoulders were vegetated with young trees, 
shrubs and weeds, providing a more open habitat for 
the yucca plants to grow and spread. The senior author 
was notified of the find, and he was able to relate the 
fact that characteristic larval “frass chimneys” should 
be present within nearby yucca plants if the skippers 
had been established at the site prior to the observa- 
tion. Subsequently, three such structures were located 
in yucca plants along the roadside. Later that day, the 
senior author confirmed the identification and pre- 


pared the collected specimen, which serves as a voucher 
for not only a new Kentucky record, but also a north- 
central extension of the known range of M. yuccae. 

On 13 April 2002 the second author and Loran D. 
Gibson visited the site. Gibson reported “20 to 25 of 
the skippers flying about and easy to approach.” More 
photos of the food plants and skippers were taken 
(Figs. 1, 2). The third author returned to the site on 26 
April 2002 and found three individuals still present. 
One was a female that appeared to be searching for 
host plants on which to lay eggs. The other two were 
males, one being very worn. 

On 18 April 2002, the second author visited the site 
again, and observed a fresh female ovipositing on a 
yucca plant, and eggs on the plant. A male and another 
individual, quite worn, were observed at this site. He 
drove along several roads in the area and found four 
additional sites with evidence of M. yuccae’s presence. 
At one, there was an unopened larval “chimney” asso- 
ciated with a Yucca plant (Fig. 3). At the second, he 


found one live pupa in a broken-off “chimney,” and 
one female ovipositing on different plants. At a third 
site he observed three unopened “chimneys,” plus 10 
eggs on yucca leaves. At the final site he found two 
“chimneys.” All sites are within a few miles of each 
other in Calloway County, Kentucky. 

The Yucca Giant-Skipper has not been published as 
occurring in Illinois (U.S.G.S. state lists) or Missouri 
(J. R. Heitzman pers. com.). Leroy GC. Koehn (pers. 
com.) reported that he has found colonies in western 
Tennessee, but these data have not been formally pub- 
lished. 

These colonies represent the northern-most known 
occurrence of a breeding colony of M. yuccae along 
the Mississippi River. On the East Coast it extends 
northward to southeastern Virginia at about the same 
latitude as the Calloway County site; and farther west 
it is found in Kansas and westward also at about the 
same latitude (Opler & Malikul 1998). 

Interestingly, Yucca has been so widely introduced 
and adventive populations have spread so readily be- 
yond it’s historic range that it is not completely clear 
how far north natural populations of this plant occur 
(J. N. N. Campbell pers. com.). According to Medley 
(1993), the species is not native to Kentucky. However, 
with populations of the plant seemingly well estab- 


Journal of the Lepidopterists’ Society 
56(4), 2002, 292-295 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


lished, it appears that the Yucca Giant-Skipper may 
now be established along with its host plant. These 
small colonies are vulnerable to habitat destruction, 
over collecting and other dangers. We hope to investi- 
gate protection of these sites and to seek other 
colonies in the area. 


We thank Loran D. Gibson for the use of his photographs of the 
live Yucca Giant Skipper. 


LITERATURE CITED 


MEDLEY, M. 1993. An annotated catalog of the known or reported 
vascular flora of Kentucky. Ph.D. Dissertation, University of 
Louisville, Kentucky. 2595 pp. 

OPLER, P. & V. MALIKUL. 1998. A field guide to the butterflies of 
eastern North America. Houghton-Mifflin, Boston. 486 pp. 


CHARLES V. COVELL JR., Dept. of Biology, University 
of Louisville, Louisville, Kentucky 40292-0001, USA; 
WILLIAM R. BLACK JR., 201 Friedman Ave., Paducah, 
Kentucky 42001-4744, USA; AND BRAINARD L. 
PALMER-BALL JR., Kentucky State Nature Preserves 
Commission, 801 Schenkel Lane, Frankfort, Kentucky 
40601-1403, USA 


Received for publication 1 September 2001; revised and accepted 10 
July 2002. 


FEMALE NORTH AMERICAN EVERES HUBNER, [1819] AND THE IDENTITY OF 
LYCAENA SISSONA W. G. WRIGHT, 1905 (LYCAENIDAE) 


Additional key words: genitalia, identification, subspecies. 


Two similar species, Everes amyntula (Boisduval, 
1852) and Everes comyntas (Godart, [1824]) (Ly- 
caenidae: Polyommatinae), are broadly sympatric and 
may fly together in parts of the western United States 
and adjacent Canada (Scott 1986, Stanford & Opler 
1993, Layberry et al. 1998, Guppy & Shepard 2001). 
The former is widespread while E. comyntas appar- 
ently occurs in scattered populations west of the Rocky 
Mountains, but, at times, the latter may be the only or 
most abundant species at some locales. Although a 
number of wing traits have been implicated as useful 
in separating these species (e.g., Klots 1951, Downey 
1975, Fisher 1981, Pyle 1981, Opler 1999), individual, 
seasonal, and geographical variation confound their 
identity. This variation has not been thoroughly inves- 
tigated and it may be that superficial differences will 
have to be elaborated at the local level. Males have 


demonstratively different genitalia, most readily seen 
in the shape of the uncus (Bethune-Baker 1913, John- 
son 1972, Dornfeld 1980, Fisher 1981, Scott 1986, 
Guppy & Shepard 2001). Females generally have not 
been distinguished except by association with males 
and superficial characters of apparently limited value. 
One character that has not been mentioned, but may 
useful in separating females of the two species, is in 
the submargin of the dorsal hindwing. All female E. 
amyntula examined from Arizona, California, Col- 
orado, Nevada, Oregon, and Utah (n = 265) had a row 
of usually several pale submarginal macules. Everes 
comyntas from Arizona and California (n = 54), how- 
ever, lacks these macules except on apparent short-day 
phenotypes having considerable dorsal blue (e.g., 
Field 1938, Shapiro 1974a). It thus seems that individ- 
uals without pale submarginal macules on the dorsal 


VOLUME 56, NUMBER 4 


hindwing are E. comyntas; those with macules could 
be either species (e.g., see Howe 1975, wherein all 
three possibilities are illustrated). Over 100 female 
Everes (76 E. amyntula, 42 E. comyntas) from the 
United States and northern Mexico have been dis- 
sected and a difference was detected between the gen- 
italia of taxa commonly placed within E. comyntas and 
E. amyntula. The genital plate (sterigma) of E. comyn- 
tas is small and rounded (Figs. 1-12), while that of E. 
amyntula is comparatively larger, more triangular, and 
some (Figs. 17, 18, 21, 22) exhibit a notch at its poste- 
rior edge (Figs. 13-24). These exhibit some variation 
within both species, locally (e.g., Figs 1-4; 19-22) and 
perhaps geographically, but their overall gestalt re- 
mains, including that of an E. comyntas from Veracruz, 
Mexico. 

Wright (1905) illustrated, named, and briefly de- 
scribed Lycaena sissona based upon a single female 
from “Sisson, Cal.” (now the city of Mt. Shasta);. The 
holotype, spread with the ventral surface upward and 
housed at the California Academy of Sciences (Tilden 
1975), has the following labels: small, white, handwrit- 
ten - / 400 /; red, printed - / HOLOTYPE °/ Lycaena / 
sissona Wright / Det. J. W. Tilden 1975 /; white, 
printed and handwritten - / W. G. WRIGHT / Plesio- 
type No. 400 / Illustrated in his / Butterflies of / the 
West Coast / Calif. Acad. Sci. Coll. /; white, printed 
and handwritten - / California Academy / of Sciences / 
Type / No. 4319 /; white, printed and handwritten - / 
Genitalic Vial / GTA - 12243 /. The specimen appears 
to be in the same condition as when it was originally 
photographed (Wright 1905). 

Lycaena sissona has largely been treated as a syn- 
onym of Everes comyntas comyntas (Comstock 1927, 
McDunnough 1938, dos Passos 1964, Miller & Brown 
1981, 1983), but was placed as a synonym of E. amyn- 
tula amyntula “based upon Wright’s illustration” by 
Ferris (1989). As far as can be determined, the type 
has not heretofore been critically examined. Wright's 
(1905) illustration is of the ventral surface and, as 
noted above, the only potentially useful characteristic 
to separate females of the two species is on the dorsal 
hindwing. The dorsal surface of the type is uniformly 
brown except for a vague orange macule proximad to a 
vague black marginal spot in hindwing cell 
CuA,—CuA,. The absence of submarginal macules and 
the configuration of its genital plate (Fig. 6) clearly 
identifies the specimen as an E. comyntas. 

Western E. comyntas, specifically those in Califor- 
nia, have not been elaborated. Dornfeld (1980), Em- 
mel et al. (1998), and Guppy and Shepard (2001) have 
all considered E. comyntas comyntas to embrace 
populations in the western United States and south- 


293 


c 
© 


© 


a 
Ses 


Fics. 1-12. Dorsal view of the genital plate of female Everes 
comyntas (numbers following data are GTA genitalic vial numbers). 
1-4. CA: Sutter Co.; Sutter bypass, Hwy. 20, E of Meridian, 29 May 
1989 (11541, 11542, 11543, 11221), 5. CA: Colusa Co.; Ca 20 at Ca 
45, W of Meridian, 29 May 1989 (11544), 6. holotype of Lycaena 
sissona (12243), 7. CT: Hartford Co.; New Britain (7129), 8. MS: 
Nachez Trace, 8 mi. S of Tupelo, 13 June 1972 (7139), 9. MN: Itasca 
Co.; Deer River, 29 June 1971 (7130), 10. WI: Iron Co.; Rt. 2, 29 
June 1971 (11219), 11. AZ: Santa Cruz Co.; Sycamore Canyon, 22 
June 1983 (11551), 12. AZ: Santa Cruz Co.; Sycamore Canyon, 9 
July 1980 (11552). 


western Canada. These have a paler and grayer venter 
than that of populations from the eastern United 
States and thus the white halos around the black discal 
macules are less distinct. The submarginal row of mac- 
ules on the ventral forewing is less complete and less 
well-defined on individuals from California as are the 


Fics. 13-24. Dorsal view of the genital plate of female Everes 


amyntula (numbers following data are GTA genitalic vial numbers). 
13. NV: Mineral Co.; White Mts., road to Sugarloaf, 5.1 mi. S of 
Montgomery Pass, 1 July 1991 (11212), 14. NV: Nye Co.; Toiyabe 
Mts., Jett Canyon, 17 May 1990 (7142), 15. NV: Clark Co.; Moapa 
Valley, California Wash, 18 June 2000 (11181), 16. CA: Nevada Co.; 
S Fork of Yuba River, N of Nevada City, 13 May 1985 (11546), 17. 
CO: Garfield Co.; White River Plateau, Coffee Pot Springs, 10100’, 
27 June 1963 (11353), 18. NV: Elko Co.; Jarbidge Mts., Bear Creek 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


marginal macules on the ventral hindwing. Also on the 
ventral hindwing, the orange macule or macules to- 
wards the tornus are notably paler in California, not 
bright orange. These characters conform with the 
holotype of Lycaena sissona and indicate that popula- 
tions in California and perhaps elsewhere in western 
North America should be treated as a recognizable 
subspecies, Everes comyntas sissona (W. G. Wright, 
1905), new combination. 

In comparison with Californian populations of E. 
amyntula amyntula, E. comyntas sissona has a more 
rounded forewing termen (this appears as a useful char- 
acter throughout the distribution of both species, con- 
tra Downey 1975), has less dorsal blue (usually without, 
except on the short-day form), the ventral hindwing or- 
ange is usually more prominent, and the ventral 
forewing macules are usually further from the submar- 
ginal macules and form a straighter row (usually more 
sinuate on E. amyntula). The male has a broader black 
margin on the dorsal forewing (usually very thin on E. 
amyntula) and usually has at least one orange macule 
on the dorsal hindwing (usually absent on E. amyntula). 
The female of E. comyntas has no submarginal pattern 
on the dorsal hindwing (again, except on the short-day 
form); these are nearly always prominent on E. amyn- 
tula. Most male E comyntas from throughout the 
species’ distribution have a thin line of black scales at 
the distal end of the forewing discal cell whereas this 
was not seen on examined E. amyntula. Females of 
both species often have a similar and sometimes 
broader black mark in the discal cell. This is most read- 
ily seen on extensively blue individuals. 

In California, E. comyntas apparently mostly inhab- 
its lowlands, frequently along ditches and other water- 
ways (Opler & Langston 1968, Shapiro 1974a, 1974b, 
but see Garth & Tilden 1963); E. amyntula is largely 
montane (Comstock 1927, Emmel & Emmel 1962, 
Shapiro et al. 1979). In Oregon, the two species fly 
sympatrically and synchronically at several sites in the 
Coast Range (fide A. D. Warren). Everes comyntas has 
a long flight season and may have two to five broods 
annually (Opler & Langston 1968, Shapiro 1974a) 
whereas E. amyntula are apparently univoltine or bi- 


ae 

Summit, 7 Aug. 1980 (7160), 19. NV: White Pine Co.; Schell Creek 
Range, Timber Creek, 5.6 mi. E of Nv 486, 15 July 1981 (7157), 20. 
NV: White Pine Co.; Schell Creek Range, Berry Creek, 2.3-5 mi. E 
of Nv 486, 16 July 1981 (11555), 21. NV: White Pine Co.; Snake 
Range, Baker Creek Campground, 16 July 1980 (7159), 22. NV: 
White Pine Co.; Steptoe Valley, Warm Springs, 24 June 1987 (7154), 
23. NV: Washoe Co.; Carson Range, Tahoe Meadows, 10 July 1990 
(11214), 24. NV: Carson City; Carson Range, Nv 28, 2.0 mi. S of 
Washoe Co. line, 1 July 1985 (7128). 


VOLUME 56, NUMBER 4 


voltine (Emmel & Emmel 1962, 1973, Shapiro et al. 
1979). Both species use a variety of legumes 
(Fabaceae) as larval hostplants (Emmel & Emmel 
1962, 1973, Shapiro 1974a, 1974b, Shapiro et al. 
1979), including alien taxa (Shapiro 2002). Whether 
there are any hostplant preferences for either species, 
except those imposed ecologically, has not been re- 
ported in California. In Colorado, E. comyntas uses 
larval hostplants occupying more mesic sites than 
those of E. amyntula (Scott 1992). 


I thank Norman D. Penny at the California Academy of Sciences 
for loaning and permitting dissection of the type of Lycaena sissona. 
Chuck Hageman and Sterling O. Mattoon graciously led me to 
populations of E. comyntas in California and allowed examination of 
specimens in their collections. Andrew D. Warren read a draft of the 
manuscript and made useful comments. 


LITERATURE CITED 


BETHUNE-BAKER, G. T. 1913. Everes comyntas and amyntula 
(Lepid.). Ent. News 24:97-103. 

Comstock, J. A. 1927. Butterflies of California. Los Angeles: publ. 
by author. 334 pp. 

DornFELD, E. J. 1980. The Butterflies of Oregon. Forest Grove, 
OR: Timber Press. 276 pp. 

DOS Passos, C. F. 1964. A synonymic list of the Nearctic Rhopalo- 
cera. Lepid. Soc. Mem. 1:1-145. 

Downey, J. C. 1975. Genus Everes Hiibner, pp. 350-351 in W. H. 
Howe (ed.), The Butterflies of North America. Garden City, 
New York: Doubleday. 633 pp. 

EMMEL, J. F., 7. C. EMMEL & S. O. Marroon. 1998. A checklist of 
the butterflies and skippers of California, pp. 825-836 in T. 
C. Emmel (ed.), Systematics of Western North American 
Butterflies. Gainesville, Florida: Mariposa Press. 878 pp. 

EMMEL, T. C. & J. F. EMMEL. 1962. Ecological studies of Rhopalo- 
cera in a high Sierran community—Donner Pass, California. 
I. Butterfly associations and distributional factors. J. Lepid. 
Soe. 16:23-44. 

EMMEL, T. C. & J. F. EMMEL. 1973. The butterflies of southern 
California. Nat. Hist. Mus., Los Angeles Co., Sci. Series 
26:1-139. 

Ferris, C. D. (ed.). 1989. Supplement to: A catalogue/checklist of 
the butterflies of America north of Mexico. Lepid. Soc. 
Mem. 3:1-103. 

FIELD, W. D. 1938. New forms and subspecies of North American 
Libytheidae and Lycaenidae. J. Kansas Ent. Soc. 11:124-133. 

FIisHER, M. S. 1981. Subfamily Polyommatinae Swainson, 1827. 
Pp. 200-223 in C. D. Ferris & F. M. Brown (eds.), Butter- 
flies of the Rocky Mountain States. Norman: Univ. Okla- 
homa Press. 442 pp. 

GarTH, J. S. & J. W. TILDEN. 1963. Yosemite butterflies. J. Res. 
Lepid. 2:1-96. 

Guppy, C. S. & J. H. SHEPARD. 2001. Butterflies of British Columbia, 
including western Alberta, southem Yukon, the Alaska Pan- 
handle, Washington, northern Oregon, northem Idaho, north- 
western Montana. Vancouver: UCB Press. 414 pp. 


Howe, W. H. 1975. The butterflies of North America. Garden City, 
New York: Doubleday, 633 pp. 

JouNson, K. 1972. The butterflies of Nebraska. J. Res. Lepid. 
11:1-64. 

Kiots, A. B. 1951. A field guide to the butterflies of North 
America, east of the Great Plains. Boston: Houghton Mifflin. 
349 pp. 

LayBerry, R. A., P. W. HALL & J. D. LAFONTAINE. 1998. The but- 
terflies of Canada. Toronto: Univ. of Toronto Press. 280 pp. 

McDunnoucu, J. H. 1938. Check list of the Lepidoptera of 
Canada and the United States of America. Part 1. Macrolepi- 
doptera. Mem. So. Calif. Acad. Sci. 1:1-273. 

MILuer, L. D. & F. M. Brown. 1981. A catalogue/checklist of the 
butterflies of America north of Mexico. Lepid. Soc. Mem. 
2:1—-280. 

MILLER, L. D. & F. M. Brown. 1983. Lycaenidae, pp- 53-57 in R. 
W. Hodges (ed.), Check list of the Lepidoptera of America 
north of Mexico. London: Classey. 284 pp. 

OPLER, P. A. 1999. A field guide to Western butterflies. 2nd ed. 
Boston: Houghton Mifflin. 540 pp. 

OpLeER, P. A. & R. L. Lancston. 1968. A distributional analysis of 
the butterflies of Contra Costa County, California. J. Lepid. 
Soc. 22:89-107. 

PyLe, R. M. 1981. The Audubon Society field guide to North 
American butterflies. New York: A. A. Knopf. 916 pp. 
Scott, J. A. 1986. The Butterflies of North America, a natural his- 
tory and field guide. Stanford, California: Stanford Univ. 

Press. 583 pp. 

ScoTr, J. A. 1992. Hostplant records for butterflies and skippers 
(mostly from Colorado) 1959-1991, with new life histories 
and notes on oviposition, immatures and ecology. Papilio 
(new series) 6:1-171. 

SHAPIRO, A. M. 1974a. The butterfly fauna of the Sacramento 

Valley, California. J. Res. Lepid. 13:73-82, 115-122, 

137-148. 

. 1974b. Butterflies of the Suisun Marsh, California. J. Res. 

Lepid. 13:191—206. 

. 2002. The Californian urban butterfly fauna is dependent 

on alien plants. Diversity Distr. 8:3140. 

SHAPIRO, A. M., C. A. PALM & K. L. WersLo. 1979. The ecology 
and biogeography of the butterflies of the Trinity Alps and 
Mount Eddy, northern California. J. Res. Lepid. 18:69-152. 

STANFORD, R. E. & P. A. OPLER. 1993. Atlas of Western USA but- 
terflies, including adjacent parts of Canada and Mexico. 
Denver and Ft. Collins: publ. by authors. 275 pp. 

TILDEN, J. W. 1975. An analysis of the W. G. Wright butterfly and 
skipper plesiotypes in the collection of the California Acad- 
emy of Sciences. Occ. Pap. Calif. Acad. Sci. 118:1-44. 

Wricut, W. G. 1905. The Butterflies of the West Coast of the 
United States. San Francisco: publ. by author. 257 pp. 


GEORGE T. AUSTIN, Nevada State Museum and His- 
torical Society, 700 Twin Lakes Drive, Las Vegas, 
Nevada 89107, USA 


Received for publication 28 May 2002; revised and accepted 14 
August 2002. 


Journal of the Lepidopterists’ Society 
56(4), 2002, 296-297 


Aethes baloghi, 230 
Aethes matheri, 228 
Aethes matthewcruzi, 232 
Aethes razowskii, 226 
Aethes sexdentata, 225 
Aethes terriae, 229 
Aethes westratei, 227 
Agraulis vanillae, 193 
Alabama, 98 

Alesa amesis, 265 

algal partners, 289 
algivory, 289 

altiplano, 66 

Andaman Islands, 123 
Andes, 66 

Anisota manitobensis, 5 
Anisota stigma, 5 
Antherea andamana, 123 
Apoecilus, 112 
Arctiidae, 9, 289 
Arecaceae, 117 
Arkansas, 162 

Artemisia tridentata, 173 
Artemisia tripartita, 173 
Austin, G. T., 293 


Bacillus thuringiensis var. kurstaki, 166 
bait traps, 178 
Baldridge, C. D., 162 
Bali, 108 

bamboo feeders, 286 
Batesia, 199 

Becker, V. O., 9, 191 

Bia actorion, 117 
biogeography, 9 
biography, 234 

biology, 5, 123, 265 
Black Jr, W. R., 290 
Boloria, 180 

Borneo, 108 

boulleti species group, 66 
brain, 193 

Brassolinae, 117 

British Virgin Islands, 9 
Brou Jr., V. A., 178 

Brou, C. D., 178 

Brown Jr., K. S., 104, 117 
Bt, 166 

Bulia, 129 


bur oak, 5, 112 


bursa copulatrix, 62 

butterfly abundance, 166 

caenides group, 53 

Calhoun |. V., 98 

calling females, 5 

Callophrys vam 

captive rearing, 173 

Caria (section Apocaria) myristicaeformis, 1 


Caribbean, oD 


INDEX FOR VOLUME 56 


(new names in boldface) 


Cary, S. J., 49, 151 
caterpillar calls, 265 
Catocala, 1, 234 
Catocala atocala, 1 
cladistics, 129, 199 
Cladonia polycarpoides, 289 
Clossiana, 180 
Cochylini, 216 

collecting expedition, 151 
collections, 234 
Colombia, 53 

colonizers, 9 

coontie, 272 

copulation, 106 

Cordero, C., 106 
Coreopsis, 176 

Cossidae, 9 

Covell Jr., C. V., 290 
cycasin, 272 

cypselus group, 53 


Dalla, 53 

Dalla celsus, 59 

Dalla disconnexa, 56 
Dalla pedro, 59 

Dalla vista, 56 

Dalla wardi, 54 

Danaus plexippus, 90 
Dang, P. T., 172 

dashed Phulia pattern, 66 
deforestation, 108 
dehydration, 90 

DeVries, P. J., 45, 183, 199, 265 
dew drinking, 90 

diana fritillary, 162 
Diehl, J. K., 112 

discal spot, 176 
distribution, 5, 98, 162 
diversity, 166 

drought, 98 

Dunford, J. C., 182 


early stages, 265, 286 
Ecuador, 53 

Edwards, William H., 234 
egg, 62 

egg resorption, 62 

El Nifio, 90 

entomologist, 151 
entomophagous, 216 
Epiblema glenni, 279 
Epiblema infelix, 282 
Epiblema scudderiana, 283 
Epiblema tripartitana, 277 
Errata, 191, 298 

essayer, 49 

Eteona tisiphone, 286 
Eucosmini, 277 

Eumaeus atala, 272 
Euphorbiaceae, 129 


Eurybia, 265 
Eurybiini, 265 
Everes amyntula, 293 
Everes comyntas, 293 


Fabaceae, 129 

Ficus, 123 

Florida, 98 

Freitas, A. V. L., 104, 117, 286 
Frey, D., 90 

Fukuda, T., 193 


Gall, L. F,, 1, 234 
genealogy, 49 

genitalia, 53, 66, 293 
Geometridae, 9 
Georgia, 98 

Goldstein, P., 184 
Grote, Augustus R., 234 
Guana Island, 9 
Guenée, Achille, 234 
Guppy, C. S., 171, 180 


habitat loss, 162 

habitats, 98 

Hampton, N., 173 

harpe, 53 

hawkmoths, 178 

Hawks, D. C., 234 
Heidya salicella, 172 
Hemileuca hera hera, 173 
Hemileuca nevadensis, 112 
Hemileucinae, 112, 173 
Henne, D. C., 5, 112 
Hesperiidae, 53, 66, 98, 290 
Hesperiinae, 66 
Heteropterinae, 53 

Hill, R., 199 

history, 49, 234 

Hodges, R. W., 189 
Holland, R., 151 

host plant, 173 

host plant protection, 272 
hostplant, 98 

Howard, Winslow J., 49 
Hyatt, J. A., 188 
Hyblaeidae, 9 

Hylephila, 66 

Hylephila blancasi, 71 
Hylephila herrerai, 68 
Hylephila pallisteri, 70 
Hylephila pseudoherrerai, 69 
Hylephila rossi, 82 
Hylephila shapiroi, 76 
Hylephila tentativa, 73 
Hypoprepia fuscosa, 289 


identification, 293 
Illiberis rotundata, 62 


immature stages, 1, 104, 117, 123, 265, 286 


VOLUME 56, NUMBER 4 


immigrant, 172 
Indonesia, 108 
insect, 272 
invasion, 108 
Ithomiinae, 104 


jewler, 49 


key to species, 129 
Kondla, N. G., 171, 180 
Koshio, C., 62 

Kroutoy, V., 193 

Kudo, S., 62 


La Nina, 90 

large moths, 9 

larvae, 5 

larval diet, 289 

larval host plant, 129 
larval survivorship, 272 
Laughlin, A. C., 129 
lectotypes, 234 
Leschenaultia, 112 


life history, 1, 104, 117, 265, 286, 289 


light traps, 178 
Lithosiinae, 289 
longevity, 62 

Louisiana, 1, 178 
Lycaena sissona, 293 
Lycaenidae, 106, 272, 293 


MacNeill, C. D., 66 
Malay Archipelago, 108 
Malaysia, 108 

Manitoba, 5, 112 
mating, 106 

mating effort, 90 
Matsumoto, K., 108 
Mechanitini, 104 
Megathyminae, 290 
Megathymus yuccae, 290 
Messett, L., 90 
metalmark butterfly, 265 
Metzler, E. H., 46, 216 
Midwestern North America, 216 
Miller, S. E., 9, 191 
Miller, W. E., 172, 216 
Mississippi, 1 

monarch butterfly, 90 
Moran, M. D., 162 
morphology, 199, 265 
Moskowitz, D. P., 289 
Mount Rainier, 180 
Murray, D., 117 
mushroom body, 193 
myrmecophily, 265 


natural history, 49, 117, 151, 162, 265 


naturalist, 49, 151 
nectar plants, 162 
Neotropics, 199, 265, 286 


neotypes, 234 

neuropil, 193 

Nevada buck moth, 112 

New Mexico, 49, 151 

new species, 53, 216 
Noctuidae, 1, 9, 129, 176, 234 
nomenclature, 171, 180 
non-target butterfly, 166 
North America, 172, 216, 293 
Notodontidae, 9 

nutmeg hickory, 1 


199, 286 


oakworms, 5 

Olethreutinae, 172, 277 
olfactory lobes, 193 

oocyte, 62 

oreal (of high mountain), 66 
Oregon, 166 

Ouachita Mountain, 162 
Ozark Mountain, 162 


painted lichen moth, 289 
palatability, 9 

Palearctic, 172 
Palmer-Ball Jr., B. L., 290 
Panacea, 199 

Papilio demoleus, 108 
Papilionidae, 108 
patronym, 49 

Patterson, W. D., 176 
Peacock, J. W., 1 

Penz, C. M., 199, 265 
Phyciodes pratensis, 171 
Phyciodes pulchella, 171 
Phyciodes pulchellus, 171 
phylogeny, 199 

Physcia millegrana, 289 
plum moth, 62 

Pogue, M. G., 129 
population, 272 

Populus, 172 

prairie, 162 

Prashanth M., 123 
predators, 272 
Procridinae, 62 
Pronophiliti, 286 

Pyrgus albescens, 98 
Pyrgus communis, 98 


Reep, R. L., 193 
re-establishment, 272 
Riodinidae, 265 
Roman, R., 90 


Sabourin, M., 172, 216 
sagebrush sheep moth, 173 
Sais rosalia, 104 

Salix, 172 

Saturniidae, 5, 112, 123, 173 


Nymphalidae, 104, 117, 162, 171, 193, 


Satyrinae, 117, 286 
season, 272 

Severns, P., 166 

sexual selection, 106 
Slotten, J. R., 1 

Smith, E. M., 272 

Snow, Frances H., 151 
Solanaceae, 104 

South America, 66, 199, 265, 286 
species abundance, 166 
species richness, 166 
spermatophore, 106 
spermatophore production, 106 
Speyeria, 180 

Speyeria bishoffti, 180 
Speyeria chariclea, 180 
Speyeria diana, 162 
Speyeria mormonia, 180 
Speyeria rainieri, 180 
Speyeria titania, 180 
Sphingidae, 9, 178 
sphinx moths, 178 
Steihauser, S. R., 53 
stigma pockets, 66 
Stiriinae, 176 
subspecies, 293 

survey, 162 

systematic position, 117 
systematics, 199, 234 


Tachinidae, 112 

Tanaka, Y., 62 

taxonomy, 9, 216, 234 
threetip sagebrush, 173 
Tortricidae, 172, 216, 277 
Tortricinae, 216 
transtilla, 53 
Trentepohlia, 289 

type localities, 180 

types, 234 


underwing moths, 1 


Vargo, J. T., 216 
Veenakumari, K., 123 


Washington, 180 
West Indies, 9 
Westphal, C., 289 
Wetlands, 162 
Wright, D. J., 277 


Xanthothrix ranculi form albipuncta, 176 


Yucca filamentosa, 290 
yucca giant-skipper, 290 


Zetheriti, 286 
Zygaenidae, 62 


ERRATA 


Journal of the Lepidopterists’ Society 
56(4), 2002, 298 


EMERGENCE OF PARASITIC FLIES FROM ADULT ACTINOTE DICEUS 
(NYMPHALIDAE: ACRAEINAE) IN ECUADOR 


In the above article by Harold. F. Greeney and John O. Stireman (Journal of the Lepidopterists’ Society 
59(2):79-80), the genus name of a parasitic fly was misspelled. Where it reads “Arachidomya sp.” (pp 79 and 80) 
it should read “Arachnidomya sp.” 


Date of Issue (Vol. 56, No. 4): 9 December 2002 


*y t 


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